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Lace Bug Genera of the World, I: Introduction, Subfamily Cantacaderinae (Heteroptera: Tingidae)

RICHARD C. FROESCHNER m i

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I. Michael Heyman Secretary Smithsonian Institution SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 574

Lace Bug Genera of the World, I: Introduction, Subfamily Cantacaderinae (Heteroptera: Tingidae)

Richard C. Froeschner

SMITHSONIAN INSTITUTION PRESS Washington, D.C. 1996 ABSTRACT Froeschner, Richard C. Lace Bug Genera of the World, I: Introduction, Subfamily Cantacaderinae (Heteroptera: Tingidae). Smithsonian Contributions to Zoology, number 574,43 pages, 39 figures, 1 table, 1996.—Included is an outline of adult taxonomic anatomy, a key to the two subfamilies .Cantacaderinae Stal and Tinginae Laporte (acceptance of separate family status for the Vianaididae excludes them from this publication), a key to the two tribes of Cantacaderinae: Cantacaderini Stal and Phatnomatini Drake and Davis, and keys to the 10 genera of the tribe Cantacaderini and 26 genera of the tribe Phatnomatini. Each genus is diagnosed, depicted in a dorsal habitus drawing of its type species, and accompanied by a list of currently included species with additions and changes published subsequent to the Drake and Ruhoff (1965) catalog. Where practical at this time, a key to the species within some of the genera is offered. Fossil forms are excluded except for three fossil species cataloged in modern genera: For two species of the Cantacaderini genus Cantacader, the new genus Paleocader is proposed with the new combinations Paleocader avitus (Drake) (type species) and P. quinquecarinatus (Germar and Berendt); the Phatnomatini species Phatnoma baltica Drake is transferred to the new combination Sinalda baltica. In the tribe Cantacaderini, the following generic changes have been made since the Drake and Ruhoff (1965a) catalog: increase from five genera to 10; present transfer of the genus Carldrakeana Froeschner from the tribe Phatnomatini; description of the new genus Paleocader for two fossil species (see paragraph above); and earlier transfers into Cantacaderini from Phatnomatini of Cyperobia Bergroth, Pseudophatnoma Blote, and Stenocader Drake and Hambleton. For genera within the Cantacaderini keys to two species of Allocader Drake, three species of Carldrakeana, two species of Ceratocader Drake, and two species of Pseudophat- noma are provided herein. In the tribe Phatnomatini generic changes since the Drake and Ruhoff catalog are as follows: herein description of two new genera, Etesinalda for type and only species, E. laticosta, new species, from the Island of Sao Tome and Exulmus for Ulmus engaeus Drake and Ruhoff, type and only species, with new combination Exulmus engaeus; two genera described elsewhere by Froeschner, Carldrakeana, Distocader; eight by Pericart, Daillea, Indocader, Microcader, Phatnomella, Pseudacalypta, Pullocader, Taphnoma, Thaicader; one by Stusak, Phatnocader; three genera transferred to Cantacaderini (see above); redefinition of Gonycentrum Bergroth, with Sinalda Distant elevated from synonymy; Minitingis Barber elevated from synonymy; and Phatnoma baltica Drake herein transferred to genus Sinalda. Within the tribe Phatnomatini keys to two species of Eocader Drake and Hambleton, two species of Gonycentrum Bergroth, two species of Indocader, three species of Microcader, two species of Minitingis Barber, four species of Plesionoma Drake, four species of Taphnoma, three species of Ulmus Distant, two species of Zetekella Drake are provided herein.

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging-in-Publication Data Froeschner, Richard C. Lace bug genera of the world / Richard C. Froeschner. p. cm.—-{Smithsonian contributions to zoology ; no. 574) Includes bibliographical references (p. ) and index. Contents: I. Introduction, Subfamily Cantacaderinae (Heteroptera: Tingidae) I. Title. II. Series. QLI.S54 no. 574 [QL523.T5] 591 s-dc20 [595.7'54] 95-5558 CIP

® The paper used in this publication meets the minimum requirements of the American National Standard for Permanence of Paper for Printed Library Materials Z39.48—1984. Contents

Page Introduction 1 Outline of Taxonomic Anatomy 1 Acknowledgments 3 Family TlNGlDAE Laporte 4 Key to Subfamilies of Tingidae 4 Subfamily CANTACADERINAE Stal 4 Key to Tribes of Cantacaderinae 4 Tribe CANTACADERINI Stal 5 Key to Genera of Cantacaderini 6 Genus Allocader Drake 7 Key to Allocader Species 7 Genus Cantacader Amyot and Serville 8 Genus Carldrakeana Froeschner, new tribal assignment 9 Key to Carldrakeana Species 10 Genus Ceratocader Drake 10 Key to Ceratocader Species 10 Genus Cyperobia Bergroth 11 Genus Nectocader Drake 12 Genus Paleocader, new genus (fossil) 13 Paleocader avitus (Drake), new combination 14 Paleocader quinquecarinatus (Germar and Berendt), new combination ... 14 Genus Pseudophatnoma Blote 15 Key to Pseudophatnoma Species 16 Genus Stenocader Drake and Hambleton 16 Genus Teratocader Drake 16 Tribe PHATNOMATINI Drake and Davis 17 Key to Genera of Phatnomatini 17 Genus Alloeoderes Drake 19 Genus Angiocader Drake 20 Genus Astolophos Distant 20 Genus Cnemiandrus Distant 21 Genus Cyclotynaspis Montandon 21 Genus Daillea Pericart 22 Genus Distocader Froeschner 22 Genus Eocader Drake and Hambleton 23 Key to Eocader Species 23 Genus Etesinalda, new genus 24 Etesinalda laticosta, new species 24 Genus Exulmus, new genus 25 Genus Gonycentrum Bergroth 25 Key to Gonycentrum Species 26 Genus Indocader Pericart 26 Key to Indocader Species 27 Genus Microcader Pericart 27 Key to Microcader Species 28

in iv SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Minitingis Barber 28 Key to Minitingis Species 29 Genus Oranoma Drake 29 Genus Phdtnocader Stusak 29 Genus Phatnoma Fieber 30 Genus Phatnomella Pericart 31 Key to Phatnomella Species 32 Genus Plesionoma Drake 32 Key to Plesionoma Species 33 Genus Pseudacalypta Pericart 33 Genus Pullocader Pericart 33 Genus Sinalda Distant 34 Genus Taphnoma Pericart 35 Key to Taphnoma Species 35 Genus Thaicader Pericart 36 Genus Ulmus Distant 36 Key to Ulmus Species 37 Genus Zetekella Drake 37 Key to Zetekella Species 38 Literature Cited 39 Index 42 Lace Bug Genera of the World, I: Introduction, Subfamily Cantacaderinae (Heteroptera: Tingidae)

Richard C. Froeschner

Introduction incorporated with appropriate citations, and subsequently Effective communication, in science as in everyday activi- described taxa are added. Where practical at this time, keys to ties, depends on general acceptance of the meaning of species of many of the genera are appended. words—their definitions. One of the main aims of the present Numerous fossil tingids have been described, "more than offering is to establish certain definitions so that all students of 160" according to Bekker-Migdisova (1962:302). No effort lace bugs may have a common point of departure for their was made to include them comprehensively in this study, but because 3 fossil forms were cataloged in modern genera by studies. Admittedly, some of these presently proposed points Drake and Ruhoff (1965a) it was necessary to consider them: will need changing, but that admission is neither a new caution Cantacader avitus Drake and Cantacader quinquecarinatus nor a reason to avoid making them, for they are formulated only (Germar and Berendt) are transferred to the new genus on the evidences at hand; as more evidences accumulate, those Paleocader; Phatnoma baltica Drake is transferred to the points can be reappraised. genus Sinalda Distant. The extent of the taxon encompassing the lace bugs varies Effective use of a key demands that the user recognize the with modern authors. Drake and Davis (1960), basing their limits of keys. First, keys operate primarily by serving as a conclusions on an intensive study of the morphology of the device for eliminating forms from further consideration. group, considered it to be a family with three subfamilies: Secondly, confidence in the correctness of the name encoun- Cantacaderinae, Tinginae, and Vianaidinae. This concept was tered by following the key can be relied upon only to the degree followed in the Drake and Ruhoff (1965a) catalog. Stys and that all constituents of the fauna are represented in the key. Kerzhner (1975), without detailed explanation but possibly After keying, verification is needed. The keyed identification following Scudder's (1959) conclusion based on a study of the must be checked by comparing the specimen with a good female genitalia of the Heteroptera, treated it as a superfamily description of the critical characters or, better yet, against including two families: Tingidae (with two subfamilies, authentically identified specimens of the named species. If Cantacaderinae and Tinginae) and Vianaididae. The Stys and forms not represented in the key are thus discovered, the key Kerzhner arrangement is followed herein. The family Tingidae must be expanded to encompass them. It is with the awareness contains over 2100 species in about 300 genera. of these limitations that these keys are offered. This is the first of a series of papers to be offered as a means of identifying the world's modern lace bugs, at least to genus, by use of keys, diagnoses, and illustrations. To each genus is OUTLINE OF TAXONOMIC ANATOMY

added a list of known species citing their original proposal and, FIGURES 1-3 as an aid in tracing the history of the species (including earlier synonyms), a page reference to location in the Drake and Modern classification of the tingids is based almost wholly Ruhoff (1965a) catalog. Taxonomic changes made subsequent on the external anatomy of the adults. This practice follows the to that catalog (new combinations, synonymies, etc.) are old philosophy that morphology permits and reflects function and that proper functioning permits continued existence in a Richard C. Froeschner Department of Entomology, MRC-105, particular environment—it is this ability of the individual to National Museum of Natural History, Smithsonian Institution, maintain and reproduce its kind that is reflected in the concept Washington, D.C. 20560. of a species. SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

clypeal spine variously prolonged as a tubercle or spine, which is sometimes jugal spine decurved or incurved. Dorsally the head may have 0-9 long or frontal spine . short spines or tubercles, some arranged in pairs (one on each dorsomeaial.spine side of midline), some placed individually on the midline. The occipital spine paired spines (tubercles) consist of the jugal pair (one member hood on each jugum), the frontal pair (usually near an imaginary line paranotum connecting antennal bases), and the occipital pair (located on callus the occiput or arising a short distance forward so as to be median carina between the eyes). The single spines (which may be subdivided lateral carina into a left and a right member) on the midline of the head suprahumeral carina include the dorsomedial (on midline slightly anterior to eyes), scutellum the clypeal (arising from the clypeus), and sometimes one clavus arising between the frontals. The pattern and individual stenocostal area development of these spines are much used in tingid taxonomy. costal area The antenna may show significant generic differences in shape, subcostal area in proportion of, and in vestiture of the individual segments. Ventrally the head bears two longitudinal laminae, the discoidal area buccuiae, forming a median trough for the first labial segment; these buccuiae may extend distinctly anterior to the apex of the sutural area head where they may converge, even to become contiguous. (membrane) The length of the four-segmented labium (beak or rostrum) varies considerably: it may reach only onto the thoracic sterna or it may extend onto the abdomen as far as the genital structures. All three thoracic segments exhibit characters used in tingid taxonomy. These characters must be used with great caution

FIGURE 1.—Generalized tingid morphology, dorsal view. because, unfortunately, the shape of all three thoracic segments, their armature, and their intersegmental proportions, as well as

The foundation for understanding tingid anatomy was expanded and summarized by Drake and Davis (1960), and it is on that treatise (with elimination of the Vianaididae) that the buccula present outline of tingid morphology (see Figures 1-3) is -antennophore based. rostrum Important to interpreting tingid anatomy is the matter of prosternal lamina infraspecific differences as shown in sexual dimorphism (such as with differing ratios of certain antennal segments), and the differences shown by macropterous and brachypterous individ- mesosternal lamina uals of the same species (see below for discussion of thorax). ostiplar pore Although these matters must be considered when interpreting a and pentreme taxon or describing a new one, it is acknowledged that the metasternal lamina descriptions and interpretations must be based on the evidences metapleural flange at hand and must so stand until additional facts come to hand. stenocostal area The common name "lace bugs" reflects the lacy appearance created by a network of fine, raised lines evident on much of the costal area dorsal surface of the insect. These lines are referred to as hypocosta "veins," and the spaces enclosed by them are termed "cells" or "areolae." The head may be porrect (projecting subhorizontally in front of eyes) or variously decurved (appearing shortened in dorsal view) anterior to eyes. The 4-segmented antennae are each inserted in a socket in a variously developed modification of the side of the head capsule; these modifications are referred to as the antennophores or antennal tubercles. The antennophore may be short or long and may have the outer apical angle FIGURE 2.—Generalized tingid morphology, ventral view. NUMBER 574 the form and sculpture of the wings, may be quite different in stenocostal area brachypterous and macropterous individuals of the same species. When a taxon is known from individuals of only one wing length, there is no way to anticipate the conditions in individuals with the other wing length; such forms must be dealt with as observed until information about the other morph permits reevaluation of the separating characters. Ventrally, the three thoracic sterna each may have a pair of longitudinal carinae or laminae (that is, with cells), one on each side of the midline, forming a narrow channel in which the ostiolar pore labium reposes at rest—these are the sternal laminae and the hypocosta space between them generally is termed the rostral groove. The prothorax is variously provided with simple carinae or laminae. The pronotal disc most commonly has three longitudi- FIGURE 3.—Cantacaderquadricornis lateral view showing alignment of nal carinae (which may be expanded into laminae with the peritreme and stenocostal area. inclusion of one or more rows of cells): the median carina and one on each side of it, the lateral carinae. These carinae may be opening, allowing outflow of the fluid from the internal scent absent or they may be partially or wholly present and gland. The scent fluid is conducted from an internal, midventral accompanied by another pair, the suprahumerals consisting of chamber to the pore by a closed tube, the vestibular duct. In a short carina over the pronotal convexity each side of the some forms the vestibular duct extends laterad so far as to place lateral carinae and next to the humerus. The median carina may the pore at the edge of the hypocosta. The metapleural surface have near its anterior end or for full length an enlargement in laterad of the pore may be modified into what is called the the form of a weak to strong tectation or an inflated bulb-like peritreme. The peritreme may be very vague or may exhibit a structure termed the cyst or hood (the latter term is used variety of discrete forms. It may consist of a trough formed by especially when the inflation extends over part or all of the two close-set carinae that are parallel or diverging. In another head). In a few instances the posterior part of the median carina form, the limiting carinae are elevated and form an apically may also be elevated variously as a second tectation or bulbous closed loop near or slightly above the ventral margin of the cyst distinctly separated from the anterior one. The lateral hypocosta. For a further modification associated with the margin of the pronotum may be smoothly rounded (ecarinate), scent-gland opening and its functioning, see the discussion of defined by a simple carina, or marked by an expansion the stenocostal area in the tribe Cantacaderini. (containing one or more rows of cells) known as the paranotum The three pairs of typically heteropteran legs, but with only (plural paranota). The posterior margin of the pronotum may be two tarsal segments, sometime have modifications of taxo- more or less prominently extended backwards as a triangular or nomic use. These modifications will be pointed out where semicircular posterior projection. appropriate. The mesothorax bears the first pair of wings, the hemelytra The abdomen, whose first visible segment is morphologi- (singular hemelytron), whose modifications are much used in cally the second segment (the true first being concealed), offers tingid taxonomy. Greater elevation of certain veins of the a few surface modifications, such as grooves, projections, and hemelytron generally divides its surface into four main areas: carinae, that are used by the taxonomist. costal area, subcostal area, discoidal area, and sutural area, the latter often overlapping. These areas offer taxonomic characters ACKNOWLEDGMENTS in distinctness of delimiting veins, in relative size and shape, and in numbers of cells or rows of cells encompassed. The This project would have been prohibitively difficult had it clavus, which is a prominent part of the hemelytron in some not been for ready access to the Carl J. Drake tingid collection heteropterous families, may be distinct and exposed (as in the now on deposit in the National Museum of Natural History of subfamily Cantacaderinae) or reduced, depressed, and con- the Smithsonian Institution, Washington, D.C., where Drake cealed (as in most members of the subfamily Tinginae) by the worked for the last eight years of his life, plus the valued triangular posterior projection of the pronotum; in some of the assistance of an earlier National Science Foundation grant coleopteroid forms, the clavus is fused imperceptibly to the rest (NSF grant GB-791) that permitted study of critical materials in of the wing and the hemelytra meet in a straight line for their several European museums a number of years ago. The full length. Ventrally the hemelytron characteristically exhibits generous cooperation of the curators of those museums at that the hypocosta, a prominent carina or lamina (most frequently time is appreciated and hereby acknowledged: M. Beier, containing one or occasionally two or more rows of cells) that Naturhistorische Museum, Vienna; L. Brundin and P. Persson, demarks the inner edge of the costal area. Naturhistoriska Riksmuseet, Stockholm; J. Carayon, Museum The metathorax bears the hind wings and, on each side National d'Histoire Naturelle, Paris; WE. China, The Natural opposite the insertion of the hind legs, a single pore, the ostiolar History Museum, London; J. G6mez-Menor, Instituto Espanol SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY de Entomologfa, Madrid; W.L.V. Hackman, Zoological Mu- illustrations thanks are due to my wife, Elsie Herbold seum, Helsinki; G. Schmitz and H. Schouteden, Congo Froeschner (as EHF), to J. Pericart (as JP), Montereau, France, Museum, Tervuren; E. Tortenese, Museo Civico di Storia and to J.M. Stusak, Brno, Czech Republic; and to Y. Sohn for Naturale, Genova; S.L. Tuxen, Zoologisk Museum, Copen- readying the figures for the printer. Thanks for able help in hagen. preparing the final manuscript go to S.B. West. Finally, thanks Thanks for the loan of study materials are due to H.C. Blote for helpful manuscript reviews go to T.J. Henry, Systematic and D.C. Geijskes, Rijksmuseum van Natuurlijke Histore, Entomology Laboratory, U.S. Department of Agriculture, J. Leiden; F.M. Carpenter, Museum of Comparative Zoology, Maldonado Capriles, now retired at Cayey, Puerto Rico, to Cambridge; P. Duarte-Rodrigues, Department de Zoologia e C.W. Schaefer, The University of Connecticut, Storrs, and two Antropologia, Faculdade de Ciertcias, Lisboa; A.C. Eyles, anonymous reviewers. Department of Scientific and Industrial Research, Nelson, New Zealand; J. Pericart, Montereau, France; A. Soos, Magyar Nemzeti Muzeum, Budapest; M.D. Webb and J. Margerison- Family TINGIDAE Laporte Knight, The Natural History Museum, London, R.L. Wenzel of The family name of the lace bugs has appeared in several the Field Museum of Natural History, Chicago, and L.-Y. spellings: Tingidae, Tingideae, Tingidida, Tingididae, Tingidi- Zheng, Nankai University, Tiangin. tae, Tingidites (Laporte's original spelling), Tingitidae, etc. Special thanks go to G.E. Steyskal for kindly and freely The spelling "Tingidae" was adopted by the International using his vast knowledge of classical languages to help Commission on Zoological Nomenclature in its Opinion 143 determine the derivation of the generic names, and to C.W. (1943a:83) and confirmed in its Bulletin of Zoological Sabrosky for aid with nomenclatorial matters. For the Nomenclature (1943b: 13).

Key to Subfamilies of Tingidae

Clavus of hemelytron more weakly developed than mesocorium and depressed below its surface, usually wholly covered by strongly produced triangular posterior margin of pronotum. Visible abdominal segments I-IV fused TlNGINAE Laporte Clavus of hemelytron developed similarly to mesocorium, not depressed below its surface, seldom with more than its very base covered by weakly produced posterior margin of pronotum. Visible abdominal segments I—II only fused (not III and IV) CANTACADERINAE Stal

Subfamily CANTACADERINAE Stal the other represented there by four of its 30 species). Fossil CantacaderariaStal, 1873:116. records are available for three species, in two genera, in the Cantacaderinae.—Drake and Ruhoff, 1965a:22. Palearctic Region. Table 1 charts the distribution of modem genera by zoogeographic regions. DIAGNOSIS.—Clavus developed equally to and on the same COMMENTS.—The subfamily Cantacaderinae appears to be a plane with the mesocorium, meeting in a straight line to form a natural and valid group whose included genera fall into two distinct claval commissure with slightly thickened edges. categories characterized by Drake and Davis (1960:78) as (1) GEOGRAPHIC DISTRIBUTION.—This subfamily is fundamen- the tribe Cantacaderini, with a stenocostal area visible dorsally tally a taxon of the southern land masses (including southern (see discussion of characters under tribe Cantacaderini for Asia) of the world; of its modern members, none is known from fuller explanation of extent of stenocostal area), and (2) the North America and only two of its modem genera are found in tribe "Phatnomini" (emended to Phatnomatini by Froeschner the Palearctic Region (one genus restricted to that region and (1981:96)), lacking the stenocostal area.

Key to Tribes of Cantacaderinae

Hypocosta near base with a trough delimited by a pair of prominently elevated crossveins leading from near apex of ostiolar canal and, in most genera, this elevation of adjacent veins continues along the ventral surface of the costal area to set off a single row of cells, the stenocostal area [Figure 3] CANTACADERINI Stal Hypocosta not thus interrupted and no stenocostal area present on ventral surface of costal area PHATNOMATINI Drake and Davis NUMBER 574

TABLE 1.—Geographic distribution of modern Cantacaderinae by tribes and genera (numbers = totals of genera and species).

Taxon Neotropics Nearctic Palearctic Oriental Ethiopian Madagascaran Australian New Zealand Oceania

CANTACADERINAE (35, 135) 6,22 1,6 16,35 11,48 2,2 5,10 1. 1 5,11 CANTACADERINI (9,53) 2,2 1,6 3,11 I, 19 1, 1 4,9 1, 1 2,4 Allocader (3) - 3 Cantacader (39) - 19 2 3 Carldrakeana (3) - 2 1 Ceratocader (2) - 2 Cyperobia 1 - Nectocader (1) 1 Pseudophatnoma (2) Stenocader (1) 1 Teratocader (1) - PHATNOMATINI (26, 82) 4,20 13,24 10, 1, 1 1,1 3,7 Alloeoderes (1) - 1 - Angiocader (1) - - 1 Astolophus (1) - - 1 Cnemiandrus (1) - - 1 Cyclotynaspis (1) - 1 - Dai//ea(l) - 1 Dislocader (1) - Eocader (2) 2 Etesinalda (1) - Ex ulmus (1) Gonycentrum (2) - Indocader (2) - Microcader (3) - Minitingis (2) 2 Oranoma (1) Phatnocader (1) - 1 Phatnoma (28) 14 5 2 Phatnomella (2) - 2 Plesionoma (4) - 4 Pseudacalypta (1) - 1 Pullocader (1) 1 Si/w/da (14) - 14 Taphnoma (4) - 4 Thaicader (1) - (1)

Zetekella (2) 2

Tribe CANTACADERINI Stal the costal area for evaporation, is much more complex than the original description would suggest—it may or may not involve Cantacaderaria Stal, 1873:116. Cantacaderini.—Drake and Ruhoff, 1965a:22. a dorsal modification of the hemelytron. Morphologically it consists of a narrow trough bordered on each side by a DIAGNOSIS.—This tribe is characterized within the entire thickened and distinctly elevated vein. In its least development family Tingidae by the presence of a "stenocostal area" (see it is a trough only across the hypocosta between two thickened, discussion below), which is always visible ventrally and subbasal veins just opposite the apex of the peritreme (e.g., sometimes also dorsally. genus Carldrakeana Froeschner). In the next step of complex- COMMENTS.—Certain generalizations can be made: (1) head ity this trough across the hypocostal lamella joins with a similar never with spines (tubercles) on midline; (2) pronotum trough formed by the outermost row of cells on the ventral never with inflated cysts; (3) paranotum never reflexed surface of the costal area between the thick costal vein and the to form a mesally opening cyst; and (4) hemelytron always with thickened subcostal vein (e.g., genus Cyperobia Bergroth). The some expression of the stenocostal area (at least on the most complex expression adds a dorsally evident differentia- hypocosta). tion of the outermost row of costal area cells into a distinct row The stenocostal area, whose total structure and location that is set off by a thickened subcostal vein (e.g., Cantacader suggest it serves as a trough to convey the scent-gland fluid to Amyot and Serville). 6 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

This interfunctioning of structures arising on two body (transferred by Froeschner (1968:246)). A fourth additional segments—scent-gland peritreme on metathorax and steno- postcatalog genus, Carldrakeana Froeschner (1968:250), was costal area on wing arising from the mesothorax—must be described in Phatnomatini but is herein transferred into the under multiple gene control and hence not easily duplicated. Cantacaderini. A fifth additional genus, Paleocader (fossil), is The steps of increasing complexity of this evaporating described herein for two fossil species from European ambers: mechanism can be termed a transformation series in the sense Cantacader avitus Drake, type species, and Tingis quinquecar- of Hennig (1966:89) and permits the tribe to be adjudged inatus Germar and Berendt. monophyletic. GEOGRAPHIC DISTRIBUTION.—All geographic regions ex- As treated herein the tribe Cantacaderini contains twice the cept the Nearctic contain one or more modern members of this number of genera cataloged in it by Drake and Ruhoff (1965a). tribe. The genus Cantacader Amyot and Serville is restricted to In addition to the five genera included there, three genera have the Old World, where it has some of its member species in each been transferred from the tribe Phatnomatini, where they were of the subregions there. Of the other eight genera, two are cataloged by Drake and Ruhoff: Cyperobia Bergroth (trans- confined to the Neotropics, two occur only in the Oriental ferred by Stusak (1979:149)); Pseudophatnoma (transferred by Region, and four are known from the Australo-New Zealand Pericart (1991:49)); and Stenocader Drake and Hambleton area (see Table 1).

Key to Genera of Cantacaderini

1. Posterior margin of pronotum convexly to triangularly produced, completely covering scutellum 2 Posterior margin of scutellum medially transverse or partly concave, exposing apex or more of scutellum 4 2. Paranotum strongly explanate, margin forming long, narrow, triangular projection anterolaterally and at midength Pseudophatnoma Blote Paranotum explanate or not, never forming large triangular projections 3 3. Hemelytral outline abruptly expanded from base, forming an extremely broad costal area (15-20 row of cells wide) for its full length Teratocader Drake Hemelytral outline gradually widening from base, costal area narrower (less than 10 rows of cells wide) Cantacader Amyot and Serville 4. Paranotum with 6-8 long spines (some as long as width of an eye) extending horizontally at right angles to lateral margin Ceratocader Drake Paranotum with not more than 1 long spine, lateral margins sometimes almost serrate 5 5. Lateral margins of paranotum and costa with numerous small but distinct triangular projections appearing as irregular serrations Stenocader Drake and Hambleton Lateral margins of paranotum and costa without projections or serrations 6 6. Costal area with 1-2 rows of cells (including stenocostal row delimited by thickened vein only on ventral side) 7 Costal area with 6 or more rows of cells (including stenocostal row delimited dorsally and ventrally) 8 7. Paranotum on anterior half with 1 row of moderately large cells and deflexed, at least in part, against propleuron. Interocular area on each side of midline with a pair of fine, black, oblique sutures extending from occiput to base of supraclypeal spines Cyperobia Bergroth Paranotum for full length areolate, obliquely elevated. Interocular area of head without fine, black, oblique sutures Carldrakeana Froeschner 8. Pronotum dorsally with only 3 distinct longitudinal carinae .... Allocader Drake Pronotum dorsally with 5 distinct longitudinal carinae 9 9. Costal area abruptly and very much widened from base, with 12 or more rows of cells. Head elongate, preocular part about 5 times as long as horizontal diameter of an eye Nectocader Drake Costal area gradually widening from base, almost continuing outline of paranotum, with 5-6 rows of cells. Head short, preocular part not more than twice as long as horizontal diameter of an eye Paleocader, new genus NUMBER 574

Genus Allocader Drake Allocader leai (Hacker).—Drake and Ruhoff, 1965*23. Cantacader leai Hacker, 1928:176 [Australia]. FIGURE 4 Allocader nesiotes Drake and Ruhoff, 1962:249 [Australia]; 1965a:23. Allocader Drake, 1950:156 [type species: Cantacader leai Hacker, original designation].—Drake and Ruhoff, 1965a:22.

DIAGNOSIS.—Among the tribal members with exposed scutellum and entire (neither spined nor serrate) paranotal margins, this genus can be recognized by its tricarinate (no suprahumeral carinae) pronotal disc and its broad (4-6 rows of cells) costal area. Length 3-9 mm. GEOGRAPHIC DISTRIBUTION.—All records for this genus are for Australia and its nearby islands. ETYMOLOGY (masculine).—allos, Greek (other), plus the nondescriptive fragment -coder from the generic name Canta- cader denoting another generic type in the taxon containing Cantacader. COMMENTS.—Examination of a paratypic A. leai nymph, with the same label information as the paratypic adult at hand, shows the thorax and the abdomen to be spineless except for a posteriorly deflexed, long, stout, sharp spine on the midline of each of the first two abdominal tergites. Its cephalic spines are short and blunt as in the adult. The nymphal prothorax shows the broad (but here nonareolate) paranotal expansions typical of the genus. The basal pronotal width given as "1.10 mm" in the original description of A. nesiotes Drake and Ruhoff must be a typographical error because the accompanying illustration shows it to be equal to half the width of the combined hemelytra described as "5.20 mm." Drake and Ruhoff (1962:250) stated in the comparative note for A. nesiotes, "The cephalic spines (two pairs in front of eyes) are long as in A. leai, whereas they are short, tuberculate in the other species." Examination of pertinent type material and original descriptions reveals that it is A. cordatus (Hacker), as correctly described by Drake with the original description of the genus, which has the long cephalic spines.

List of Allocader species Allocader cordatus (Hacker). Phatnoma cordata Hacker, 1927:19 [Australia]. Allocader cordata.—Drake and Ruhoff, 1965a:22. FIGURE 4.—Allocader leai, natural length 3 mm.

Key to Allocader Species

1. Cephalic spines tapering, as long as or longer than horizontal width of an eye. Posterior margin of pronotum deeply, triangularly emarginate medially A. cordatus Cephalic spines reduced to short, blunt tubercles less than half as long as the horizontal diameter of an eye. Posterior margin of pronotum not emarginate medially 2 2. Paranotum broad, nearly half as wide as pronotal disc, nearly horizontal and distinctly areolate A. leai Paranotum less than half as wide as pronotal disc, nearly vertical and vaguely areolate A. nesiotes SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Cantacader Amyot and Serville

FIGURE 5

Cantacader Amyot and Serville, 1843:299 [type species: Piesma quadricornis Lepeletier and Serville, only included species].—Drake and Runoff, 1965a:23. DIAGNOSIS.—Cantacader differs from all other members of the tribe in the combination of the wholly hidden scutellum, lack of spine-like or angular projections on the paranotal margin, and the costal outline being gradually widened from the base (not abruptly flared). Length 3.3-4.9 mm. GEOGRAPHIC DISTRIBUTION.—This genus is restricted to the Old World, where it has representatives on all major land masses and numerous remote islands. ETYMOLOGY.—The original description states this name is from the Sanskrit words cantaca (a spine) and dri (to carry), in obvious reference to the presence of the enormous cephalic spines. The -er ending makes the name masculine. In subsequent practice the component -cader (without meaning in Greek or Latin) is often used in the formation of other generic names in the subfamily Cantacaderinae (e.g., Allocader, Ceratocader, Stenocader, etc.) and is always treated as masculine. COMMENTS.—The two fossil species cataloged in Canta- cader by Drake and Ruhoff (1965a), avitus Drake and quinquecarinatus Germar and Berendt, are unlike all other members of the genus in that they have an exposed scutellum; here they are transferred to the new genus Paleocader, where they are discussed further. Now only modern forms remain in Cantacader. The components of this genus, as herein defined, appear to form a very homogeneous group, so much so as to suggest that there may be too many species described in it. Adding to the difficulty in species separation is the existence of sexual dimorphism as well as varying degrees of brachyptery within some species; the brachyptery introduces changes in the shape and proportion of the areas of the hemelytra, changes in pronotal width, and changes in the shape of the longitudinal pronotal carinae. Until the variability of the several FIGURE 5.—Cantacader quadricornis, natural length 4.2 mm. characters can be evaluated in several species by larger series from one population, an accurate key to species can not be constructed. Cantacader amplicostatus Duarte-Rodrigues, 1981a:135 [Nigeria]. List of Cantacader Species Cantacader amydis Drake, 1960:343 [D'Entrecasteaux Islands].—Drake and Cantacader abdivitus Drake, 1950:161 [Australia].—Drake and Ruhoff, Ruhoff, 1965a:24. 1965a:23. Cantacader angulipennis Horvath, 1906:12 [Spain].—Drake and Ruhoff, Cantacader afzelii Stal, 1873:116 [Sierra Leone].—Drake and Ruhoff, 1965a:24. 1965a:23. Cantacader angustecostatus Stusak, 1979:142 [Congo Republic; Ghana]. Cantacader agilis Drake, 1951:166 [Bismarck Islands].—Drake and Ruhoff, Cantacader attenuatus Distant, 1902a:238 [South Africa].—Drake and Ruhoff, 1965*24. 1965a:24. Cantacader agilis agilis Drake, 1951:166 [Bismarck Islands]. Cantacader basilewskyi Schouteden, 1955b: 163 [Ruanda].—Drake and Ru- Cantacader agilis tricarinatus Drake. hoff, 1965a:24. Cantacader agilis var. tricarinatus Drake. 1951:167 [Bismarck Islands].— Cantacader bowmansi Schouteden, 1965a: 170 [Zaire]. Drake and Ruhoff, 1965a: 24. Cantacader claini Schouteden, 1965a: 171 [Zaire]. Cantacader allaeri Schouteden, 1965a: 169 [Zaire]. Cantacader claratis Drake, 1950:160 [Malaya].—Drake and Ruhoff, 1965a:25. NUMBER 574

Cantacader curtulus Linnavuori, 1977:6 [Yemen]. Cantacader diffidentis Drake and Poor, 1936:141 [India].—Drake and Runoff, 1965a:25. Cantacader divisus Bergroth, 1908:108 [Ethiopia].—Drake and Runoff, 1965a:25. Cantacader formosus Drake, 1950:159 [Formosa].—Drake and Runoff, 1965*25. Cantacader gerardi Schouteden. 1955b: 162 [Congo].—Drake and Runoff, 1965a:25. Cantacader hubtaerti Schouteden, 1965a: 171 [Zaire]. Cantacader ilongaensis Duarte-Rodrigues, 1982:326 [Tanzania]. Cantacader infuscatus Distant, 1903:124 [India].—Drake and Runoff, 1965a:25. Cantacader insularis Drake, 1957:399 [Reunion Island].—Drake and Runoff, 1965a:25. Cantacader japanicus Drake, 1947:225 [Japan].—Drake and Ruhoff, 1965a:26. Cantacader laratanus Drake, 1947:226 [Larat Island].—Drake and Ruhoff, 1965a:26. Cantacader laticollis Horvith, 1906:11 [Algeria].—Drake and Ruhoff, 1965a:26. Cantacader letabanus Duarte-Rodrigues, 1981b:202 [South Africa]. Cantacader lethierryi Scott, 1874:291, 443 [Japan].—Drake and Ruhoff, 1965a:26. Cantacader longicornis Duarte-Rodrigues, 1980:3 [Malawi]. Cantader nocturnis Hacker. Cantacader nocturnis Hacker, 1929:324 [Australia].—Drake and Ruhoff, 1965a:26. Cantacader quadricornis (Lepeletier and Serville).—Drake and Ruhoff, 1965a:26.. Piesma quadricornis Lepeletier and Serville, 1828:653 [Spain]. Cantacader quadricornis nubilus Horvith. Cantacader quadricornis var. nubilus Horvith, 1906:12 [Caucasus; TUrkey].—Drake and Ruhoff, 1965a:27. Cantacader quadricornis quadricornis (Lepeletier and Serville). Piesma quadricornis Lepeletier and Serville, 1828:653 [Spain]. Cantacader quadricornis quadricornis.—Horvdth, 1906:12. Cantacader quinquecostatus (Fieber).—Drake and Ruhoff, 1965a:27. Taphrostethus quinquecostatus Fieber, 1844:41 ["Ostindien"]. FIGURE 6.—Carldrakeana tindalei, natural length 2.2 mm. Cantacader schoutedeni Stusak, 1984:237 [Zaire]. Cantacader sejunctus Duarte-Rodrigues, 1987b:350 [South Africa]. Cantacader tener Bergroth, 1894:167 [Madagascar].—Drake and Ruhoff, 1965a:28. Cantacader tenuipes Stal, 1865:26 [Sierra Leone].—Drake and Ruhoff, a single row in the basal two-thirds or with only 2 rows 1965a:28. separated, above and below, by a slender, unspecialized vein) Cantacader tenuipes furtivus Drake. plus lack of serrations on the margins of paranota and costa. Cantacader tenuipes var. furtivus Drake, 1950:153 [Congo].—Drake and Length 2.1-2.6 mm. Ruhoff, 1965a:28. Cantacader tenuipes tenuipes Stal.—Drake, 1950:153. GEOGRAPHICAL DISTRIBUTION.—Carldrakeana occurs in Cantacader uniformis Distant, 1902b:353 [India].—Drake and Ruhoff, Australia (including Tasmania), New Guinea, and New 1965a:28. Zealand. Cantacader vandenplasi Schouteden, 1923:83 [Congo].—Drake and Ruhoff, ETYMOLOGY.—This name was originally stated to be a 1965a:29. patronym dedicated to Carl Drake but was given a feminine ending. Genus Carldrakeana Froeschner, new tribal assignment COMMENTS.—This taxon is that part of the genus Gonycen- trum, in the broad sense of the Drake and Ruhoff (1965a) FIGURE 6 catalog, characterized by the absence of spines or tubercles Carldrakeana Froeschner, 1968:250 [type species: Phatnoma tindalei Hacker, between the eyes. Three of the species listed there are currently original designation]. placed in this genus. DIAGNOSIS.—This genus may be recognized within the tribe Members of this genus show a partial development of the by the reduced number of rows of cells in the costal area (either stenocostal area and they therefore appear to represent the 10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY transition from the tribe Phatnomatini to the tribe Cantacade- part of the costal area belongs to C. socia, and figure 3 with the rini. The costal area of its species with a single row of cells costal area wholly uniseriate belongs to C. engista. could not show differentiation into an outer and inner part, and List of Carldrakeana Species the species with a double row do not have the vein between them thickened. Nevertheless, in all three included species the Carldrakeana engista (Drake and Ruhoff).—Froeschner, 1968:251. Gonycentrum engistum Drake and Ruhoff. 1961:127 [New Guinea]; hypocostal lamella does have the subbasal cross veins opposite 1965a:33. [Note: See "Comment" above correcting figure number in the small scent-gland opening noticeably thickened to form a original proposal.] narrow transverse trough leading to the costal area. This trough Carldrakeana socia (Drake and Runoff)-—Froeschner, 1968:251. across the hypocosta is unique to the tribe Cantacaderini and Gonycentrum socium Drake and Ruhoff, 1961:128, fig. 2 [Tasmania]; must be recognized as the beginning of the transition series 1965a:33. leading from the well-developed stenocostal area present in Cyperobia carectorum.—Woodward, 1961:156 ["subbrachypterous" data only, fig. 19]. most members of the tribe Cantacaderini, to which Carldrak- [Notes: (1) See "Comment" above correcting figure number in original eana is herein newly assigned. proposal. (2) Lack of a clear understanding of the true nature of Cyperobia Comparison of the written parts of the original descriptions carectorum Bergroth led Froeschner (1968) to assign several earlier of Carldrakeana engista (Drake and Ruhoff) and Carldrak- records of that species to C. socia. A corrected full synonymy is given above.] eana socia (Drake and Ruhoff) with their respective holotypes Carldrakeana tindalei (Hacker).—Froeschner, 1968:251. find they agree; the captions for the illustrations, however, are Phatnoma tindalei Hacker, 1928:177 [Australia]. transposed—figure 2 with the double row of cells in the apical Gonycentrum tindalei.—Drake and Ruhoff, 1965a:34.

Key to Carldrakeana Species

1. Pronotum unicarinate, median carinae well developed, lateral carinae absent.... C. engista Pronotum tricarinate, lateral carinae as well developed as median 2 2. Costal area irregularly biseriate to base C. tindalei Costal area biseriate only on apical third or less, uniseriate on basal two-thirds or more C. socia

Genus Ceratocader Drake ETYMOLOGY (masculine).—Greek keratos (hom), probably

FIGURE 7 referring to the large, thick spines on the head, plus the nondescriptive fragment -coder (masculine) from generic name Ceratocader Drake, 1950:157 [type species: Cantacader armatus Hacker, Cantacader, denoting another generic type in the taxon original designation].—Drake and Ruhoff, 1965a:29. containing Cantacader. DIAGNOSIS.—Within the tribe, this is the only genus with List of Ceratocader Species several prominent slender spines on margin of paranotum. Length 3.5-4.5 mm. Ceratocader armatus (Hacker).—Drake and Ruhoff, 1965a:29. Cantacader armatus Hacker, 1928:174 [Australia]. GEOGRAPHICAL DISTRIBUTION.—Of the two included spe- Ceratocader dentatus (Hacker).—Drake and Ruhoff, 1965JU29. cies, one is known from Australia, the other from Tasmania. Cantacader dental us Hacker, 1928:175 [Tasmania].

Key to Ceratocader Species

Lateral margin of paranotum with 7-8 acute spines directed outward, some spines longer than diameter of an eye. Costal area oblique, in no part recurved C. armatus Lateral margin of paranotum with about 6 shorter spines, all distinctly shorter than diameter of an eye. Costal area on anterior half recurved over itself. . . . C. dentatus NUMBER 574 11

Genus Cyperobia Bergroth FlGURE i.—Cype cHF FIGURE 8

Cyperobia Bergroth, 1927:673 [type species: Cyperobia carectorum Bergroth, monobasic].—Drake and Ruhoff, 1965a:31. pying the greatest part of the anterior pronotal lobe is DIAGNOSIS.—Within this tribe, Cyperobia is definable by misleading in suggesting the existence of a bladder or elevated the following combination of features: posterior margin of inflated cyst; actually there is only a slight dorsal convexity on pronotum transverse, medially weakly, angularly concave, the pronotal collar. fully exposing scutellum; and margins of paranota and costae This genus was cataloged under the tribe "Phatnomini" by simple, without spines or serrations. Length 3.7-4.3 mm. Drake and Ruhoff (1965a:31), but, as pointed out by Stusak GEOGRAPHIC DISTRIBUTION.—The single species of the (1979:149) and confirmed by examination of additional New genus is known only from New Zealand. Zealand specimens, its sole member has a strongly developed ETYMOLOGY (masculine).—Greek kyperos (Latin cyperus) stenocostal area ventrally and must be placed in the tribe (sedge), plus Greek bios (a manner of life), in reference to the Cantacaderini. Dorsally the subcostal vein is perceptibly wider early report of it being found on a sedge plant. than the cross veins, but not as markedly elevated as it is on the COMMENTS.—The original description of a "vesicle" occu- ventral surface. 12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

List of Cyperobia Species Genus Nectocader Drake Cyperobia carectorum Bergroth ["species of Tingid"].—Myers, 1922:6 FIGURE 9 [footnote only; New Zealand]; 1926:477 [ICassinia leptophylla]. Cyperobia carectorum Bergroth, 1927:674 [New Zealand].—Myers and Nectocader Drake, 1928:41 [type species: Cantacader gounellei Drake, China, 1928:380.—Woodward, 1961:154 ["macropterous" data only].— monobasic].—Drake and Runoff, 1965a:29. Froeschner, 1968:251 [part]. Carldrakeana species.—Froeschner, 1968:251 [for Woodward's, 1961, fig. DIAGNOSIS.—The combination of the partially exposed 18]. scutellum plus the very broad costal area (broader than REMARKS.—For this lone species of Cyperobia the above subcostal or discoidal areas) dilating abruptly from base listing is given in detail to complete deciphering certain early marks this genus for ready recognition within the tribe. Length confusions with Carldrakeana socia. Myers (1922:6, footnote) 6.4 mm. reported this species, without a name, from "swamp vegeta- GEOGRAPHIC DISTRIBUTION.—The lone species of the genus tion," with the remark that his specimen had been sent to Bergroth for determination. Bergroth (1927:674) described the single adult as Cyperobia carectorum and included thereunder Myers' (1926:477) comments as "possibly from Cassinia leptophylla" Through the cooperation of A. Soos and A.C. Eyles, an opportunity to examine 18 specimens of C. carectorum, including two macropterous and 16 brachypterous individuals, made it possible to establish the true identity of the species, even without seeing the single specimen on which Bergroth based his original description. Study of a brachypterous male labeled "Sedges. Gollan's V., 5-2-21," in agreement with the collecting data given in the original description, was especially interesting. Bergroth gave the single measurement of a female, so this male cannot be the holotype, unless the sex of the type was reported in error. This specimen is in the Entomology Division of the Department of Scientific and Industrial Research in Nelson, New Zealand, and would be available for neotype designation if one were needed. Froeschner's (1968) misinterpretation of Bergroth's use of "vesicle" led him to erroneously conclude that Woodward's (1961:155) figure 18 was not of this species. However, his figure 19 of a brachypterous individual was a misidentification of a specimen of Carldrakeana socia, and in the text of that paper all references to individuals showing reduced wing development apparently reflect this latter species. Both parts of the scientific name associate this species with sedges. The generic name translates freely as a dweller on sedges, and the specific name suggests that this insect utilizes plants of the sedge genus Carex as a host. However, labels on specimens examined (see list below) reported two plant associations, one a sedge, the other a member of the family Asteraceae. The following New Zealand specimens examined for this study were mostly brachypterous, only two from the Mt. Harper series were macropterous: Gollan's Valley, 2 Feb 1921, sedges; Mt. Jolloes, 26 Oct 1962, A.C. Eyles, Celmisia spectabilis; Dashwood Pass, 5 Mar 1962, Cassinia leptophylla; ML Harper, 14 Feb 1962, J.I. Townsend, 4000 feet, Celmisia spectabilis; ML Hutt, Canterbury, 1 Feb 1962, J.I. Townsend; Karori, 8 Mar 1924. FIGURE 9.—Nectocader gounellei, natural length 6.4 mm. NUMBER 574 13 is known only from Brazil. ETYMOLOGY (masculine).—Latin necto (knit), plus the nondescriptive fragment, -cader (treated as maculine) from the generic name Cantacader, in combination denoting the knitted or lace-like appearance of a genus in the same group as the genus Cantacader. COMMENTS.—This genus appears to be very close to Teratocader Drake because of its great size, length of head, number of cephalic spines, angular projection of posterior margin of pronotum at level of clavo-corial suture, shape and relative width of costal area, and long slender legs. The most significant difference between the two lies in the greater prolongation of the pronotal midline of Teratocader, where that structure completely covers the scutellum (exposed in Nec- tocader). Except for that point and the great geographic gap between the ranges of the two genera, the two species involved could be placed in one genus with little difficulty. However, if any importance is to be attached to the posterior prolongations of the pronotum as part of a progressive development within the Cantacaderinae, this one character must weigh importantly. If one of the two species (each known from but one specimen) is found to be mislabeled, the matter must be reconsidered. The original description and subsequent comments about the costal areas of Nectocader gounellei (Drake) and the very similar Teratocader magnificus (Drake) being uniseriate resulted from misinterpreting the stenocostal area for the costal area. Actually the costal area (laterad of the hypocostal lamella) is quite broad and contains approximately 10-15 rows of small cells, in addition to the single row in the stenocostal area.

List of Nectocader Species Nectocader gounellei (Drake).—Drake and Runoff, 1965a:29. E||F Cantacader gounellei Drake, 1923:81 [Brazil]. FIGURE 10.—Paleocader avitus, natural length 3.9 mm. Genus Paleocoder new genus (fossil)

FIGURE 10 avitus (Drake). Bucculae slightly surpassing, and almost DIAGNOSIS.—The smooth, unarmed paranotal margins, the contiguous beyond apex of clypeus (not visible in original four long slender cephalic spines, plus the exposed scutellum figures of P. quinquecarinatus). Labium reaching to or beyond combine to mark this fossil genus within the tribe. All basal third of abdomen. comments given herein concerning P. quinquecarinatus were Pronotum without inflated cysts; anterior margin sinuately derived from its original description and its accompanying transverse. Disc with five distinctly elevated longitudinal illustration. See discussion of that species below. carinae: median and lateral pairs reaching anterior pronotal CHARACTERS.—Length 3.0-3.9 mm. Macropterous; heme- lytral axes subparallel; rounded apices slightly separated. margin, interrupted at calli; suprahumeral pair somewhat Head wide, width across eyes greater than length, preocular curved, restricted to interhumeral convexity. Paranotum mod- part about one and one-half times as long as horizontal erately explanate, in P. avitus biseriate anteriorly, uniseriate diameter of an eye, with four long tapering spines (jugals and around humerus (not discernible in original figure of P. frontals). Eyes somewhat bulbous, almost half as wide as quinquecarinatus). Posterior margin weakly convex, exposing interocular width. Antenna slender, cylindrical, segment I scutellum. shorter than interocular width, in P. avitus (Drake) slightly Hemelytra with costal margins diverging from bases, longer than II and in P. quinquecarinatus (Germar and Berendt) greatest combined width near midlength. Discoidal area one-third as long as II; IV missing from unique specimen of P. reaching apical two-fifths of hemelytron, five to six cells wide, 14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY with one or two distinctly elevated cross veins. Subcostal area Ruhoff, 1965a:24. weakly oblique, with 5-7 rows of cells and 1 or 2 distinctly Paleocader quinquecarinatus (Germar and Berendt) [new combination]. Tingis quinquecarinata Germar and Berendt, 1856:23 [fossil! in Prussian elevated crossveins. Costal area in P. avitus with five rows of amber]. cells in addition to stenocostal row (see discussion below for Cantacader quinquecarinatus.—Drake and Ruhoff, 1965a:27. costal area of P. quinquecarinatus). Hypocosta probably uniseriate. Paleocader avitus (Drake), new combination The peritreme and other ventral features were not adequately described or reliably illustrated for either of the species Drake's original placement of avitus in Cantacader was (obscured by opaque cloud in amber around holotype of P. logical because the specimen possessed the following features avitus; see discussion of P. quinquecarinatus below). How- in common with modern members of that genus: presence of a ever, judging from the well-developed stenocostal area dorsally distinct claval commissure; four long, tapering, cephalic on the holotype of "Cantacader avitus," the peritreme might spines; the presence of a stenocostal area dorsally; and several be distinctly formed and elevated to be a part of the other proper pronotal and hemelytral features. Examination of ostiolar-stenocostal system, at least in that species. the holotype confirmed the original illustration showing an TYPE SPECIES.—Cantacader avitus Drake, present designa- exposed scutellum (not covered by an expansion of the tion. posterior pronotal margin). In this instance, the exposed GEOGRAPHIC DISTRIBUTION.—Both included species were condition must be the ancestral condition that existed before the described from European amber: Paleocader avitus from posterior expansion and hence the species could not belong to "Baltic amber," generally considered to be from Oligocene a later-appearing genus, Cantacader, with a well-marked deposits on the island of Rugen off the Baltic Coast of posterior expansion. Here the unspecialized structure of the Germany; and P. quinquecarinatus from "Prussian amber," pronotum and the earlier geologic occurrence are used to justify which is also dated as of Oligocene origin. its assignment to a new genus. ETYMOLOGY (masculine).—Greek, palaios (ancient), plus -cader (masculine) from the generic name Cantacader denot- ing another generic type in the taxon containing Cantacader. Paleocader quinquecarinatus (Germar and Berendt), COMMENTS.—The virtually unexpanded posterior margin of new combination the pronotum of this otherwise Cantacader-likc genus suggests Tingis quinquecarinatus is transferred herein to this genus, that Paleocader represents an earlier time in the line leading to even though no specimens were available for study, and the the modern genus Cantacader wherein the scutellum is always original presentation described or illustrated features that raise covered by the pronotum. An alternate hypothesis, that severe doubts as to the wisdom of this assignment. The dorsal Paleocader may represent a derived state resulting from loss of habitus, illustrated without a head, is so typically that of a the expanded posterior margin of the pronotum, is argued cantacaderine that an effort is made to explain away certain against by the time factor because no true Cantacader is known contradicting evidences. to have occurred in an earlier geological time. The lack of a stenocostal area in the sketch, and unmentioned Scudder (1890:359) placed the fossil species "Tingis quin- in the text, may be explained by many of the cells along the quecarinata" Germar and Berendt in his new genus Eotingis, which he proposed for his new fossil species, Eotingis costal margin being shown as half circles, suggesting either a antennata, from Miocene rocks in North America. That reflexion of that margin, and hence obscuring of a stenocostal association of these two lace bugs in the same genus area, or, equally likely, a degree of "artistic license" in omitting undoubtedly was based on the fact that both were fossil forms some of the smaller details and overlooking the restricted outer rather than on comparison of their morphology. The dorsal row of cells. The taxonomic value of the stenocostal area was sketch accompanying the original description of T. quinquecar- not known at that time. inata, in spite of lacking a head, shows the habitus and many of The sketch of a ventral view with head attached presents a the structural features of a Cantacader-Wke Cantacaderinae; in number and magnitude of much more serious problems, which, contrast, the figure accompanying the original description off. if confirmed by study of the actual specimen, could require antennata presents quite different fades showing the critical important revision in thinking about the classification of this structures of members of the subfamily Tinginae. These facts part of the Tingidae. The head is unlike that of any other led Drake and Ruhoff (1965a) to keep the two species in two Cantacaderinae because it is very short (scarcely exceeding separate genera, each in a different subfamily. Plans are to treat anterior margin of eye), possibly foreshortened due to resting at Eotingis as a member of the Tinginae in a subsequent part of an angle in the amber, and, unlike any other Tingidae, it has a this series. very long second antennal segment. These two features of the A fuller discussion of the two species included herein is head, plus its absence in the dorsal view illustration, lead to the given after listing below. suspicion that the head was erroneously associated in recon- List of Paleocader Species structing the specimen while drawing. The following characters Paleocader avitus (Drake) [new combination]. in the ventral sketch appear unexplainable (except for Cantacader avitus Drake, 1950:161 [fossil! in Baltic amber].—Drake and erroneous depiction) and very much unlike other Tingidae: NUMBER 574 15

FIGURE 11.—Pseudophatnoma corniculata, natural length 6.7 mm.

relative widths of pro- and pterothorax; the prosternum the latero-anterior angle of the paranotum, coupled with the completely surrounding the anterior coxae; and the absence of strongly and convexly projecting posterior pronotal margin an indication of a hypocosta. covering the scutellum, identify this genus within the subfam- ily Cantacaderinae. Length = 6.7 mm. Genus Pseudophatnoma Blote GEOGRAPHIC DISTRIBUTION.—Reported distribution for this

FIGURE 11 genus includes the Riau Archipelago, off the tip of Malay Peninsula, and the island of Borneo. Pseudophatnoma B16te, 1945:78 [type species: Pseudophatnoma corniculata B16te, monobasic].—Drake and Runoff, 1965a:40. ETYMOLOGY (neuter).—pseudos, Greek (false), plus generic Froeschnerocader Pericart, 1986:245 [type species: Froeschnerocader denti- name Phatnoma, suggesting a general similarity to Phatnoma collis Pericart. monobasic. Synonymized by Pericart, 1991:40]. but emphasizing that it is not the same as that genus. DIAGNOSIS.—The forward-directed spine-like extensions of COMMENTS.—Due to the lack of specimens for study and the 16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY fact that the strongly reflexed costal area concealed the own examination of that type confirmed those actions. stenocostal area in the illustration accompanying the original description, this genus was cataloged in the tribe "Phatnomini" List of Pseudophatnoma Species by Drake and Ruhoff (1965a:40), a placement that misled Pseudophatnoma corniculata BlOte, 1945:78 [Riau Archipelago].—Drake and Pericart into describing his new genus, Froeschnerocader. Ruhoff, 1965a:40. Pericart's personal examination of Blote's type discovered the Pseudophatnoma denticoUis.—Pericart, 1991:50. synonymy and the necessity for the transfer to this tribe. My Froeschnerocader denticoUis Pericart, 1986:246 [SabahJ. Key to Pseudophatnoma Species

Anterior margin of pronotum transverse, not forming an angle above base of head P. corniculata Anterior margin of pronotum medially forming a prominent angle projecting almost to imaginary line connecting anterior margins of eyes P. denticoUis

DIAGNOSIS.—The marginally serrated appearance, resulting from the presence of widely spaced, irregular, subtriangular projections along the edges of the paranota and costae, identifies this genus within the subfamily. Length 3.3-3.8 mm. GEOGRAPHIC DISTRIBUTION.—The single species of this genus is cataloged only for Chile. ETYMOLOGY (masculine).—stenos, Greek (narrow), plus the nondescriptive -cader (masculine) from the generic name Cantacader denoting another generic type in the taxon containing Cantacader. COMMENT.—The stenocostal area is not evident dorsally, but it is clearly delimited ventrally, where it is set off by a strongly elevated vein. List of Stenocader Species Stenocader tingidoides (Spinola).—Drake and Ruhoff, 1965a:40. Piesma tingidoides Spinola, 1852:200 [Chile].

Genus Teratocader Drake

FIGURE 13 Teratocader Drake, 1950:158 [type species: Cantacader magnificus Drake, only included species].—Drake and Ruhoff, 1965a: 29. DIAGNOSIS.—Within the subfamily, members of this genus can be recognized by the combination of the very broad costal area abruptly convexly dilated from base, plus the posterior margin of the pronotum angularly expanded to cover the scutellum. Length 8.6 mm. GEOGRAPHIC DISTRIBUTION.—The sole species is known only from the Malay Peninsula. ETYMOLOGY (masculine).—teratos, Greek (monster), plus the nondescriptive -cader (masculine) from the generic name Cantacader denoting another generic type in the taxon containing Cantacader. The unusually large size of this species FIGURE 12.—Stenocader tingidoides, natural length 3.6 mm. undoubtedly suggested using the above prefix. COMMENTS.—See remarks in "Comments" under the genus Genus Stenocader Drake and Hambleton Nectocader about misinterpretations of stenocostal area for costal area and general similarities between the two genera. FIGURE 12 list of Teratocader Species Stenocader Drake and Hambleton, 1944:120 [type species: Piesma tingidoides Teratocader magnificus (Drake).—Drake and Ruhoff, 1965a:30. Spinola. only included species].—Drake and Ruhoff. 1965a:40. Cantacader magnificus Drake, 1923:83 [Malaya]. NUMBER 574 17

DIAGNOSIS.—Within the subfamily this tribe is marked by the total absence (even from the hypocosta) of the stenocostal area. CHARACTERS.—The characters of this tribe are much as given in the above description for the subfamily Cantacaderi- nae. Within that description and, unlike the three restrictions described above for the tribe Cantacaderini, members of the Phatnomatini (1) never have an indication of a stenocostal area, (2) may have spines or tubercles on the midline of the head, and (3) may have the paranotum strongly reflexed back over itself (in one genus—Angiocader Drake). The hypocosta, unless described otherwise, can be assumed to be 1-seriate. GEOGRAPHIC DISTRIBUTION.—This is a tribe of the Southern Hemisphere; none of its members enter the Nearctic or Palearctic regions. Phatnoma Fieber is the most widespread genus, occurring throughout most of the southern hemisphere. Gonycentrum Bergroth is the only other genus reportedly not restricted to one zoogeographic area; it is known from Asia and Africa. Each of the remaining 24 genera is restricted to a single zoogeographic region: 13 in the Oriental; 10 in the Ethiopian; 4 in the Neotropics; and 3 in Oceania. COMMENTS.—Several generic changes from the Drake and Ruhoff (1965a) cataloging of this tribe will be noticed: Cyperobia Bergroth, Pseudophatnoma Blote, and Stenocader Drake and Hambleton are no longer listed herein, having been transferred to the tribe Cantacaderini earlier in the present paper. Postcatalog publications restored Minitingis Barber to generic status and added Daillea Pericart, Distocader Froesch- ner, Indocader Pericart, Microcader Pericart, Phatnocader Stusak, Phatnomella Pericart, Pseudacalypta Pericart, Pullo- cader Pericart, Taphnoma Pericart, and Thaicader Pericart. In addition, two new genera are described herein: Etesinalda for the new and only species, E. laticosta, and Exulmus for its type EJIr and only species, Ulmus engaeus Drake and Ruhoff. The nominate genus name, Phatnoma, as pointed out to me several years ago by George Steyskal, is based on the neuter Tribe PHATNOMATIN1 Drake and Davis Greek noun phatnoma (genitive phatnomatos, stem phat- Phatnomini Drake and Davis, 1960:68.—Drake and Runoff, 1965a:30. nomat-); thus, the tribal name derived from it must be Phatnomatini.—Froeschner, 1981:96. Phatnomatini as first used in publication by Froeschner (1981).

Key to Genera of Phatnomatini

Interocular area of head distinctly depressed, eyes appearing obliquely elevated above it. Costal margin a carina basally, thence abruptly expanded a short distance from base Eocader Drake and Hambleton Interocular area of head not depressed, eyes not appearing elevated. Costal margin forming a continuous line to base 2 2. Head with a dorsomedial spine or tubercle (not to be confused with spine near base of clypeus) 3 Head without a dorsomedial spine or tubercle 18 3. Head with a pair of jugal spines 4 Head with no jugal spines 16 18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

4. Head with a pair of interocular or occipital spines or tubercles (sometimes greatly reduced and requiring careful examination) 5 Head without interocular or occipital spines or tubercles 14 5. Paranotum and costal area very broad, in large part curved upward and back over themselves Angiocader Drake Paranotum and costal area narrow or broad, never curved upward and back over themselves 6 6. Head with an elongate median spine between frontals (anterior to dorsomedial) Ulmus Distant Head without median spine between frontals 7 7. Paranotum extremely narrow, on posterior lobe becoming obsolete around humerus Etesinalda new genus Paranotum with 1 or more rows of cells around humerus 8 8. Paranotum with lateral margin convex, straight, or slightly sinuate, with no lobular or spined projections 9 Paranotum with lateral margin forming rounded, angulate, or spined projections, at least at anterior end 11 9. Peritreme not forming an elevated, apically closed loop Sinalda Distant Peritreme strongly developed as an apically closed loop 10 10. Paranotum 3-seriate, greatest width across paranota distinctly anterior to humeri. Labium not reaching metacoxae Daillea Pericart Paranotum 1- or 2-seriate, greatest width across paranota at level of humeri. Labium reaching or surpassing posterior margin of metasternum Microcader Pericart 11. Pronotal disc 3-carinate 12 Pronotal disc 1-carinate 13 12. Posterior paranotal lobe narrow, 2-seriate Phatnomella Pericart Posterior paranotal lobe very wide, 5-6-seriate Plesionoma Drake 13. Shape ovate, width almost two-thirds of length. Paranotum with 3 or more rows of cells Indocader Pericart Shape elongate, width slightly less than half of length. Paranotum 1 seriate, except on anterior third Phatnocader Stusak 14. Pronotal disc 3-carinate Distocader Froeschner Pronotal disc 1-carinate 15 15. Form (brachypters only known) broadly oval, width more than half of length. Paranotum with lateral margin irregular, anteriorly forming an angle, thence abruptly dilated to form truncate 3-seriate lobe Exulmus new genus Form (macropters only known) elongate, width less than half of length. Paranotum with lateral margin simply convex or forming a large angle opposite humerus Taphnoma Pericart 16. Buccula, in profile, strongly triangular, 5-6 cells high posteriorly, tapering to 1 cell anteriorly Pseudacalypta Pericart Buccula subequal in height for full length, with 1 row of cells 17 17. Pronotum across lobular expansions of paranota wider than width of combined hemelytra. Pronotal disc with no longitudinal carina AUoeoderes Drake Pronotum without lobular expansions of paranota, narrower than combined width of hemelytra. Pronotal disc with percurrent median carinae . . . Thaicader Pericart 18. Spines on dorsum of head nearly as long as or longer than length of head Oranoma Drake Spines on dorsum of head not more than half as long as head 19 19. Abdomen ventrally on basal half or more with a distinctly impressed mediolongi- tudinal groove. Paranotum narrowest opposite humerus, thence widened cephalad to 4 or more rows of cells Phatnoma Fieber Abdomen ventrally without mediolongitudinal groove. Paranotum anteriorly not or only slightly (1-2 cells) widened 20 NUMBER 574 19

20. Head with a clypeal spine 21 Head without a clypeal spine 25 21. Occipital spines nearly or quite as long as horizontal diameter of an eye .... 22 Occipital spines absent or much shorter than an eye 23 22. Paranotal margin with 3 acutely angled projections Minitingis Barber Paranotal margin simple (without angular projections) Gonycentrum Bergroth 23. Ostiolar pore a conspicuous hole whose rim overlaps marginal vein of hypocosta Zetekella Drake Ostiolar pore confused with pleural surface, not evident 24 24. Pronotal disc 3-carinate (macropters only known). Eyes normal size, width of 1 slightly more than one-third of interocular space Pullocader Pericart Pronotal disc 1 -carinate (brachypters only known). Eyes much reduced, width of 1 about one-fifth as wide as interocular space Cyclotynaspis Montandon 25. Pronotal disc 1 -carinate, with median carina becoming evanescent in posterior third. Without paranota Astolophos Distant Pronotal disc 3-seriate, median carina percurrent, lateral carinae evident only on posterior and anterior slopes of posterior lobe. Paranotum with a single row of distinct cells continuing around humerus Cnemiandrus Distant

Genus Alloeoderes Drake

FIGURE 14

Alloeoderes Drake, 1961:115 [type species: Alloeoderes davao Drake, monobasic].—Drake and Ruhoff, 1965a:30.

DIAGNOSIS.—The members of this genus can be recognized by either of two features unique within the tribe: the broad lateral expansion of the paranota making the pronotal width more than three times that of head; or the complete absence of longitudinal carinae on the pronotal disc. Length 2.0 mm. GEOGRAPHIC DISTRIBUTION.—The single specimen was collected on Mindanao in the Philippine Islands. ETYMOLOGY (feminine).—alloio, Greek (another kind), plus dere, Greek (neck), plus adjectival ending es; doubtlessly given in reference to the unusual collar-like appearance of the uniquely expanded side margin of the paranotum. COMMENTS.—The above description was prepared wholly from the original description because no specimens were available for examination; the holotype being the only known specimen. In describing this genus and its only included species, Drake (1961) commented on the abdomen being withdrawn well into the cavity of the convex hemelytra.

List of Alloeoderes Species

Alloeoderes davao Drake, 1961:116 [Philippine Islands].—Drake and Ruhoff, 1965a:30.

FIGURE 14 (left).—Alloeoderes davao, natural length 2.0 mm. 20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Astolophos Distant

FIGURE 16

Astolophos Distant, 1904b:428 [type species: Astolophos capitatus Distant, monobasic].—Drake and Ruhoff, 1965a:30.

DIAGNOSIS.—In combination, the lack of cephalic spines on the dorsum of the head anterior to the eyes plus the lack of areolate paranota on posterior pronotal lobe distinguish this genus from all others in the tribe. Length 3.1-3.5 mm. GEOGRAPHIC DISTRIBUTION.—This is one of several tingid genera whose lone species occurs only in South Africa. ETYMOLOGY (masculine).—The origin and application are unclear. G.E. Steyskal suggested the possible derivation from asty, Greek (city), and lophos Greek (crest).

List of Astolophos Species Astolophos capitatus Distant, 1904b:429 [South Africa].—Drake and Ruhoff, 1965a:30.

FIGURE 15.—Angiocader obesa, natural length 3.0 mm.

Genus Angiocader Drake

FIGURE 15

Angiocader Drake, 1950:159 [type species: Phatnoma obesa Distant, mono- basic].—Drake and Ruhoff, 1965a:30. DIAGNOSIS.—This genus is well marked within the tribe by the shape of the paranota and costae, which are strongly recurved upward and back over themselves. Length 3.0 mm. GEOGRAPHIC DISTRIBUTION.—The lone species is known only from Cape Colony in South Africa. ETYMOLOGY (masculine).—angeion, Greek (a receptacle), plus -coder from the name Cantacader, denoting another generic type in the taxon containing Cantacader—probably given in reference to the dish-like shape produced by the elevated margins. COMMENTS.—It is worth noting that this is the only genus in the subfamily Cantacaderinae exhibiting the recurved paranota and costal area, a development that is exhibited by several genera in the subfamily Tinginae. List of Angiocader Species Angiocader obesus (Distant).—Drake and Ruhoff, 1965a:30. Phatnoma obesa Distant, 19O2a:239 [South Africa]. FIGURE 16.—Astolophos capitatus. natural length 3.1 mm. NUMBER 574 21

Genus Cnemiandrus Distant The original card containing the type series (in The Natural History Museum, London) once held six specimens and now FIGURE 17 has only four, with the original first and fourth missing. Here Cnemiandrus Distant, 1902a;239 [type species: Cnemiandrus typicus Distant, the second specimen from the right end (fifth from the left in monobasic].—Drake and Runoff, 1965a:31. the original series) is chosen lectotype—it shows the heaviest DIAGNOSIS.—The absence of cephalic spines or tubercles sclerotization; the other three, now considered paralectotypes, anterior to the eyes, combined with the presence of a distinct, are quite teneral. though narrow, uniseriate paranotum around humeral angle, List of Cnemiandrus Species permit separation of this genus from all others in the tribe. Cnemiandrus typicus Distant, 1902a: 240 [South Africa].—Drake and Runoff, Length 3.0 mm. 1965a:31. GEOGRAPHIC DISTRIBUTION.—The only species of the genus is restricted to South Africa. Genus Cyclotynaspis Montandon ETYMOLOGY (masculine).—Origin of this name is obscure. FIGURE 18 G.E. Steyskal suggested the Greek knemia (variant), might suggest "another kind" but that andros, Greek (man), seems Cyclotynaspis Montandon, 1892:265 [type species: Cyclotynaspis acalyptoides Montandon, monobasic].—Drake and Runoff, 1965a:31. inappropriate for a tiny insect, plus masculine ending -us. COMMENTS.—The only species in this genus is noteworthy DIAGNOSIS.—Among the Phatnomatini genera with a long, for having numerous, well-separated, stellate hairs on all stout spine on the clypeus between the jugal spines, this genus thoracic pleurae and on the venter of the first abdominal is uniquely marked by greatly reduced eyes and the obscure segment. peritreme for the scent-gland opening. Length 1.7-1.8 mm.

EttF

FIGURE 17.—Cnemiandrus typicus, natural length 3.0 mm. FIGURE 18.—Cyclotynaspis acalyptoides, natural length 1.7 mm. 22 SMITHSON1AN CONTRIBUTIONS TO ZOOLOGY

GEOGRAPHIC DISTRIBUTION.—The lone species in this DIAGNOSIS.—Among the genera with 8-9 head spines (or genus has been reported only from Singapore. tubercles) and the occipital pair shorter than the diameter of an ETYMOLOGY (feminine).—kykl, Greek (circle), plus aspis, eye, Daillea is defined by the paranotum being flat and Greek (shield), referring to the rounded shield-like shape of the continued with rows of cells around the humerus and its outer hemelytra. margin simple (no spines or prominent angles), the labium not reaching the posterior coxae, and the peritreme forming an COMMENTS.—The holotype was the only known specimen apically closed loop. Length 1.9 mm. of this genus until specimens were discovered in 1967 in forest GEOGRAPHIC DISTRIBUTION.—The lone species is known floor litter in Singapore by D.H. Murphy of the University of only from Sabah. Singapore (see Froeschner, 1968:246). ETYMOLOGY (feminine).—A patronym for Lucien Daille, a List of Cyclotynaspis Species French coleopterist. Original description treated this name as Cyclotynaspis acalyptoides Montandon, 1892:265 [Singapore].—Drake and feminine by ending the species name with the letter "a." Runoff, 1965a:31. COMMENTS.—None.

Genus Daillea Pericart List of Daillea Species Daillea tricostata Pericart, 1991:48, figs. 19-21 [Sabah]. FIGURE 19 Daillea Pericart, 1991:47 [type species: Daillea tricostata Pericart, monoba- Genus Distocader Froeschner sic]. FIGURE 20 Distocader Froeschner, 1968:248, 249 [type species: Malula charieis Drake and Ruhoff, monobasic].

FHF

FIGURE 19.—Daillea tricostata. natural length 1.9 mm. FIGURE 20.—Distocader charieis, natural length 2.1 mm. NUMBER 574 23

DIAGNOSIS.—Among those genera of the tribe Phatnomatini bearing a dorsomedial spine or tubercle, Distocader can be recognized by the absence of occipitals plus no lobular or spine-like projection on the free margin of the oblique paranotum. Length 2.1 mm. GEOGRAPHIC DISTRIBUTION.—The single specimen of the only species was described from New Guinea. ETYMOLOGY (masculine).—Derivation originally stated to be the Latin disto (be different), plus nondescriptive -cader, fragment from Cantacader, indicating another genus in the taxon containing that genus. COMMENTS.—The species Malala chareies, on the basis of the taxonomically important cephalic spines, is generically distinct from the type species of the genus Malala and so could not follow it into the genus Gonycentrum—the above genus had to be proposed to contain it.

List of Distocader Species Distocader charieis (Drake and Runoff).—Froeschner, 1968:250. Malala charieis Drake and Ruhoff. 1965b:244 [New Guinea].

Genus Eocader Drake and Hambleton

FIGURE 21

Eocader Drake and Hambleton, 1934:436 [type species: Eocader vergrandis Drake and Hambleton, monobasic].—Drake and Ruhoff, 1965a:31. Montea Bruner, 1940:246 [type species: Montea bouclei Brunei", monobasic. Synonymized by Monte, 1942:104]. DIAGNOSIS.—Within the tribe Phatnomatini, this genus can be recognized by either of the characters stated in the first couplet of the above key. Length 2.0-2.3 mm. GEOGRAPHIC DISTRIBUTION.—Two species are known, one each from Brazil and Cuba. ETYMOLOGY (masculine).—eos, Greek (early), plus nonde- scriptive -cader to indicate another generic type in the taxon containing the genus Cantacader. The significance of the name is unclear. FIGURE 21.—Eocader vergrandis, natural length 2.1 mm. COMMENTS.—The shape of the vertex is especially notewor- thy. The area between the eyes of all the adults and the single nymph studied is distinctly lower than the eyes and the convex recognized to genus by the unique, depressed vertex and the anteocular part of the head; this depression extends obliquely supraclypeal and clypeal tubercles. In general appearance the forward and outward as a sulcus in front of each eye. The nymph is elongate oval, without spines, and closely covered general impression created by this modification is that the with tiny, flat, stellate vestiture. vertex failed to develop fully. List of Eocader Species In the original description of the synonym Montea, Bruner Eocader bouclei (Bruner).—Drake and Ruhoff, 1965a:32. pointed out its closeness to Eocader, but held the two Montea bouclei Bruner, 1940:246 [Cuba]. distinguishable on the 1-carinate pronotum of Eocader and the Eocader vergrandis Drake and Hambleton, 1934:436 [Brazil].—Drake and 3-carinate pronotum of Montea. Later Monte (1942:104) Ruhoff, 1965a:32. reported a series of E. vergrandis from Rio de Janeiro in which some individuals were 3-carinate and some 1-carinate, thus Key to Eocader Species eliminating the only important separating feature. The bra- chypterous specimens are 1-carinate with the posterior pronotal Paranotum on anterior lobe strongly, subtriangularly explanate, lobe concave, and the macropterous specimens are 3-carinate its width there greater than transverse width of an eye with the posterior pronotal lobe convex, the lateral carinae E. vergrandis extending from the calli to the posterior margin of the Paranotum not explanate on anterior lobe, its width there pronotum. distinctly less than transverse diameter of an eye The single available nymph of E. bouclei can readily be E. bouclei 24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

incurved, but not contiguous, beyond apex of clypeus. Labium extremely long, reaching base of genital structures. Antennal segment I slightly longer than II, I plus II about as long as IV. Pronotum without elevated cyst. Anterior margin concave, laterally weakly convex behind eyes. Median carina percurrent, areolate only between calli virtually absent. Paranotum extremely narrow, with distinct cells only at level of calli. Posterior margin transverse, faintly convex medially. Scutel- lum exposed. Hemelytra conjointly convex, areas well defined by elevated veins, with 2 or 3 distinctly elevated cross veins in discoidal and subcostal areas. Discoidal area reaching apical fourth of hemelytron. Subcostal area 5-7 cells wide (except at extreme base). Costal margin abruptly expanding from very base, thence broadly, convexly curved. Costal area 5-6 cells wide. Peritreme elevated, an apically closed loop. Sternal laminae present on all 3 sterna, lateral carinae virtually straight, widely separated and somewhat outcurved at posterior apex. Abdomen broadly but shallowly impressed along midventral line of pregenital segments. TYPE OF GENUS.—Etesinalda laticosta, new species, herein designated. GEOGRAPHIC DISTRIBUTION.—The holotype, the only known specimen, came from the island of Sao Tome off the west coast of Africa. ETYMOLOGY (feminine).—etes, Greek (a neighbor), plus the generic name Sinalda Distant, alluding to the fact that this new genus is a taxonomic as well as a geographic neighbor of Sinalda. COMMENTS.—This genus appears to be quite close to Sinalda and is undoubtedly an insular offshoot thereof. The alliance is marked by the tribal features plus the identical pattern of cephalic tubercles and the broadly auriculate peritreme apex. The important differences lie in the extreme reduction in the paranotal width, the virtual lack of lateral carinae, the extremely broad costal area that is abruptly and FIGURE 22.—Etesinalda laticosta, new species, natural length 4.0 mm. broadly expanded from its very base, and the very prominent postmedian tectation along the lateral discoidal vein. The latter tectation has no counterpart in Sinalda or other Genus Etesinalda, new genus Phatnomatini genera except that it is sometimes weakly shown in Phatnoma. FIGURE 22

DIAGNOSIS.—The presence of dorsomedial, jugals, frontals, and occipitals, the virtual lack of paranotum except opposite Etesinalda laticosta, new species the calli, plus the extremely broad costal area expanding DIAGNOSIS.—As the only member of the genus, laticosta abruptly from base combine to mark this genus as distinct must be diagnosed by the generic characters. within the tribe Phatnomatini. CHARACTERS.—Length 4.0 mm. Shining, yellow brown CHARACTERS.—Length 4 mm; macropterous; hemelytra with antennal IV, eyes, scutellum, and apex of elytral tectation axes subparallel, apices broadly overlapping. virtually black; remainder of surface with weak fuscous Head with 9 tubercles (jugals, frontals, occipitals, dorsome- markings. dial, and 1 each at midlength and apex of clypeus). Eye about TYPE.—Holotype female; Lagoa Amelia, Sao Tome, 9 Sep half as wide as interocular space. Bucculae surpassing and 1949, 5200 feet; The Nat. Hist. Mus., London. NUMBER 574 25

DISTRIBUTION.—Island of Sao Tome, off west coast of Africa. HOST PLANT.—Unrecorded. COMMENT.—The specific name was suggested by the very broad costal area.

Genus Exulmus, new genus FIGURE 23 DIAGNOSIS.—Among the genera of Phatnomatini with jugals, frontals, dorsomedial, and no occipitals, this genus may be recognized by the broad paranotum with a strong marginal sinuation subapically. CHARACTERS.—Length 2.4-2.7 mm; brachypterous; heme- lytra meeting in a straight line for full length, axes weakly converging posteriorly. Head with 6 long, erect, spines (jugals, frontals, dorsome- dial, and 1 on clypeus). Eye about half as wide as interocular space. Bucculae surpassing and parallel beyond apex of clypeus. Labium reaching second abdominal sternite. Antenna slender, cylindrical: segment I almost 3 times as long as II, and as long as IV. Pronotum without inflated or elevated cyst. Anterior margin shallowly concave. Disc 1-carinate, median carina percurrent, slightly projecting beyond anterior margin. Paranotum broad, abruptly constricted in anterior third. Posterior margin trans- verse, feebly 2-sinuate. Scutellum exposed. Hemelytra meeting in a straight line for full length, conjointly convex, costal margin horizontal. Areas clearly marked by elevated veins, discoidal area reaching apical third Ej

genus, one is known only from India and Ceylon (Fieber's original locality given as "Ostindien" probably should be translated as eastern India); the other was described from the African country of Chad. ETYMOLOGY (neuter).—gony, Greek (a joint or node), plus centrum, Latin (100), of unclear application. COMMENTS.—Froeschner (1968:246-248) showed the traditional Gonycentrum of the Drake and Ruhoff (1965a: 32- 34) catalog to be composed of three morphological groups, each occurring in a different zoogeographic region. In that concept Gonycentrum reverted to a monobasic Indian genus; Distant's name Sinalda was resurrected for the African species; the species of the Australian region were placed in the new genus Carldrakeana; and the type species of Malala, M. bulliens, was shown to be conspecific with the type species of Gonycentrum, and thus was synonymized under the latter name. Later, however, Linnavuori (1977:6) described G. sinuaticolle, which was transferred to Sinalda by Duarte- Rodrigues (1981b:207). The latter author (1978:13) described an African species in Gonycentrum in this restricted sense. Then Jing (1980:400, 403) described Malala tuberculum, which is herein transferred to the genus Taphnoma Pericart. Two specimens from the Distant collection in The Natural History Museum, London, were examined, and the one bearing a label "Malala bulliens Dist. Type; Distant Coll. 1911-383," apparently in Distant's handwriting, is herein designated the lectotype of that species. It also bears the following additional labels: (a) [a red circled label reading] "Type. H. T.;" (b) Peredeniya, Ceylon, 6-09, 2406; the other specimen is to be considered a paralectotype. List of Gonycentrum Species Gonycentrum chadense Duarte-Rodriques, 1978:13 [Chad]. Gonycentrum coronatum (Fieber).—Drake and Ruhoff, 1965a:33. Teleia coronata Fieber, 1844:56 ["Ostindien"]. Malala bulliens Distant, 1910:101 [Ceylon].—Drake and Ruhoff, 1965a:34 FIGURE 24.—Gonycentrum coronation, natural length 2.3 mm. [synonymized by Froeschner, 1968:248].

Key to Gonycentrum Species

Collar and hemelytron basad of claval midlength white. Subcostal area 4-seriate G. chadense Collar and hemelytra uniformly colored, without white areas described above. Subcostal area irregularly 3-seriate G. coronatum

Genus Indocader Pericart closed loop. Length 1.9-2.4 mm.

FIGURE 25 ETYMOLOGY (masculine).—Indo, for India the country of origin, plus nondescriptive -cader, from Cantacader, indicat- Indocader Pericart, 1981:596 [type species: Indocader loebli Pericart, ing another generic type in the taxon containing that genus. monobasic]. GEOGRAPHIC DISTRIBUTION.—India and Nepal. DIAGNOSIS.—Among the Phatnomatini with 8-9 spines COMMENTS.—The type series of both included species were or tubercles on the head, Indocader is recognized by the taken at altitudes of 1500 to 3100 meters. occipitals being shorter than an eye, paranotum wide, List of Indocader Species 3-seriate and with outer margin distinctly undulate or Indocader besucheti Pericart, 1983:593 [Nepal]. 2-lobed, pronotal disc 1-carinate, and peritreme an apically Indocader loebli Pericart, 1981:597 [India]. NUMBER 574 27

FIGURE 25.—Indocader loebli, natural length 2.3 mm.

Key to Indocader Species Paranotum along lateral margin deeply incised, forming two unequal lobes. Costal area distinctly widening in basal third /. loebli Paranotum along lateral margin very slightly concave, not forming prominent lobes. Costal area of equal width virtually to base I. besucheti

Genus Microcader Pericart forming an apically closed loop. Length 1.6-1.8 mm. GEOGRAPHIC DISTRIBUTION.—India; Thailand. FIGURE 26 ETYMOLOGY (masculine).—micro, Greek (small), plus Microcader Pericart, 1981:601 [type species: Microcader unicostatus Pericart, -cader, nondescript!ve, from genus Cantacader, to indicate yet original designation]. another genus in the taxon containing that genus, in reference to DIAGNOSIS.—Among the genera of Phatnomatini with 8-9 the small size. head spines or tubercles and the occipitals being shorter than an List of Microcader Species eye, Microcader can be defined by the paranotum being seriate Microcader thai Pericart, 1991:37 [Thailand]. around the humerus with outer margin simple (no spines or Microcader unicostatus Pericart, 1981:601 [India]. lobes), the labium surpassing posterior coxae, and peritreme Microcader variegatus Pericart, 1981:603 [India]. 28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Microcader Species

1. Costal area 2-seriate Af. thai Costal area 1-seriate 2 2. Pronotum 1-carinate (median only) Af. unicostatus Pronotum 3-carinate (median and 2 laterals) Af. variegatus

dorsomedial spine or tubercle, Minitingis can be recognized by its seven short head spines (only jugals, frontals, and elongate occipitals, plus one on clypeus) plus the short, acute angula- tions on the paranotal margin. Length 1.7-2.6 mm. GEOGRAPHIC DISTRIBUTION.—The two species of the genus are known only from the Bahamas and Greater Antilles in the West Indies. ETYMOLOGY (feminine).—minimus, Latin (least), plus tingis, from the generic name Tingis Fabricius, to indicate this was one of the smallest lace bugs known at that time. COMMENTS.—Considering only the three species cataloged by Drake and Ruhoff under the generic name Zetekella Drake,

y

FIGURE 26.—Microcader unicostatus, natural length 1.8 mm.

Genus Minitingis Barber

FIGURE 27

Minitingis Barber, 1954:7 [type species: Minitingis minusculus Barber, monobasic]. Zetekella.—Drake and Ruhoff, 1965a:41 [part]. [Note: This taxon was cataloged as a junior synonym of Zetekella Drake by Drake and Ruhoff (see above) but was returned to generic status by Froeschner (1968:251).]

DIAGNOSIS.—Among the genera of Phatnomatini having no FIGURE 27.—Minitingis minusculus. natural length 1.7 mm. NUMBER 574 29 the tendency was to follow them in treating Minitingis as a Froeschner (1968:251) to resurrect Barber's genus for the West synonym of that genus. However, the appearance of a second Indies forms. West Indies species agreeing with minusculus Barber in the List of Minitingis Species narrow form, head armature, long labium, paranotal develop- ment, and grooved abdomen created a distinct morphological Minitingis elsae Froeschner, 1968:253 [Jamaica]. Minitingis minusculus Barber, 1954:7 [Bahamas].—Froeschner, 1968:251, pattern of West Indies versus continental America species. This 253. pattern appears to have true zoogeographic significance and led Zetekellaminuscula—Drake and Runoff, 1965a:41.

Key to Minitingis Species

Costa with alternate, conspicuous black and white quadrate marks, and with 4 rows of areolae M. elsae Costa without alternate black and white marks, and with 2 rows of areolae M. minusculus

Genus Oranoma Drake

FIGURE 28

Oranoma Drake, 1951:165 [type species: Oranoma biroi Drake, monobasic].— Drake and Ruhoff, 1965a: 34. DIAGNOSIS.—Among those genera of the tribe Phatnomatini with the forward-projecting spiniform prolongation of the antero-Iateral angle of the paranotum, this one may be characterized as lacking a dorsomedial but having five other head spines nearly or quite as long as the head (frontal, occipitals, and one on clypeus). Length 2.5 mm. GEOGRAPHIC DISTRIBUTION.—The single specimen of this species was from New Guinea. ETYMOLOGY (neuter).—ora, Latin (rim or edge), possibly referring to the broad costal margin, plus -noma (meaningless), and probably derived from the generic name Phatnoma Fieber (and so neuter) to indicate a degree of relationship thereto. COMMENTS.—The original generic description listed the 1-carinate pronotal disc as a feature for separating this genus from Phatnoma; however, the existence of two taxa of Phatnoma with 1-carinate pronotal disc (P. agviates and P. varians unicarinata) negates that condition as a distinguishing generic feature. The important cephalic spine development (five on Oranoma, seven on Phatnoma) remains an effective separating feature for these two genera. List of Oranoma Species Oranoma biroi Drake, 1951:166 [New Guinea].—Drake and Ruhoff, 1965a:34.

Genus Phatnocader Stusak

FIGURE 29 Phatnocader Stusak, 1976:13 [type species: Phatnocader froeschneri Stusak, monobasic]. DIAGNOSIS.—Among the genera with 8-9 head spines or tubercles (occipital shorter than an eye) this one is recognized FIGURE 28.—Oranoma biroi, macropterous, natural length 2.5 mm. by the combination of t he paranotum being mostly 1-seriate 30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tpr

FIGURE 29.—Phatnocader froeschneri, natural length 2.1 mm. FIGURE 30.—Phatnoma laciniatum, natural length 4.0 mm. around the humenis and forming a weak lobe opposite the end of the collar, plus the peritreme forming an apically closed DIAGNOSIS.—Among the Phatnomatini genera with no loop. Length 2.0-2.1 mm. dorsomedial spine and the anterolateral paranotal angle GEOGRAPHIC DISTRIBUTION.—Known only from Java. projecting as a distinct angle or spiniform process, Phatnoma ETYMOLOGY (masculine).—Fragments of two generic differs in having seven spines (jugals, frontals, occipitals, and names, Phatnoma Fieber and Cantacader, combined to one on clypeus) on head. Length 3.0-4.5 mm. indicate the present genus belongs to the subfamily Cantacader- GEOGRAPHIC DISTRIBUTION.—This genus occurs in south- inae. ern Asia (Oriental region), Africa, Malagasy, Australia, the List of Phatnocader Species Papuan and Oceanic regions, and the Neotropics. Phatnocader froeschneri Stusak, 1976:14 [Java]. ETYMOLOGY (neuter).—The definition of the Greek neuter noun, phatnoma (sunken panel or framed structure), aptly Genus Phatnoma Fieber describes the appearance of the dorsal surface that is broken FIGURE 30 into small areas surrounded by distinctly elevated veins. Phatnoma Fieber. 1844:57 [type species: Phatnoma laciniata Fieber, mono- Unfortunately, Fieber treated the name as feminine as basic].—Drake and Ruhoff. 1965a:35. evidenced by his use of the feminine ending in his species NUMBER 574 31

"laciniata." Subsequent authors, including Drake and Ruhoff Phatnoma maculatum Monte [emendation]. (1965a:35), followed Fieber. In the following list of species the Phatnoma maculata Monte, 1946:252 [Brazil].—Drake and Ruhoff, I965a:37. endings are thus emended. In practice the component "-noma" Phatnoma marmoratum Champion [emendation]. at the end of Phatnoma has been used in the formation of other Phatnoma marmorata Champion, 1897:3 [Panama].—Drake and Ruhoff, generic names to indicate relationship to the genus Phatnoma 1965a:37. (neuter) rather than using the Greek -oma, which signifies "an Phatnoma maynei Schouteden, 1916:289 [Congo].—Drake and Ruhoff, 1965a: 37. eating sore"—as in "carcinoma." Phatnoma ovatum Champion [emendation]. COMMENTS.—Phatnoma, like Cantacader, is unusual Phatnoma ovata Champion, 1897:4 [Guatemala].—Drake and Ruhoff, among the Cantacaderinae in occupying a range that extends 1965a:38. into several geographic regions of the world; the other genera Phatnoma pacifica Kirkaldy, 1908:363 [Fiji Islands].—Drake and Ruhoff, 1965a:38. of the subfamily have their species restricted to one or Phatnoma takasago Takeya, 1933:32 [Taiwan].—Drake and Ruhoff, 1965a:38. only two such regions. Most of the species of Phatnoma Phatnoma togulare Drake [emendation]. are similar in structure and variability and therefore are Phatnoma togularis Drake, 1950:154 [India].—Drake and Ruhoff, 1965a: difficult to separate; because only about half of the 27 38. Phatnoma tonkinana Drake and Maa, 1955:1 [Viet Nam].—Drake and Ruhoff, described species are available for study, no key to them is 1965a:38. offered at this time. Phatnoma trinidadana Drake, 1948:21 [Trinidad].—Drake and Ruhoff, 1965a:38. The specimen of P. baltica described from "Baltic amber" is Phatnoma uichancoi Drake, 1950:155 [New Guinea].—Drake and Ruhoff, clearly not a member of this genus. In spite of Drake's 1965a:38. statement (1950:153) with the original description that the Phatnoma varians Drake, 1922:352 [French Guiana].—Drake and Ruhoff, difference in cephalic spines "does not seem to be of generic 1965a:39. Phatnoma varians var. unicarinatum Drake [emendation]. importance," subsequent studies found these structures in the Phatnoma varians var. unicarinata Drake, 1922:353 [French Guiana].— Cantacaderinae to be very valuable at the generic level; Drake and Ruhoff, 1965a:39. therefore, the pattern of cephalic armature in baltica causes its Phatnoma varians var. varians Drake. present transfer to the genus Sinai da, which Froeschner Phatnoma varians Drake, 1922 [see above]. Phatnoma varians var. varians.—Drake, 1922:353. (1968:248) resurrected from synonymy. Phatnoma veridicum Drake and Maa [emendation]. List of Phatnoma Species Phatnoma veridica Drake and Maa, 1955:2 [Palau].—Drake and Ruhoff, 1965a:39. Phatnoma agviates Drake and Ruhoff, 1961:130 [Solomon Islands]; 1965a:35. Phatnoma vernoniae Drake and Hambleton. Phatnoma ainatum Drake and Ruhoff [emendation]. Phatnoma veroniae [sic] Drake and Hambleton, 1938:51 [Brazil]. Phatnoma ainata Drake and Ruhoff, 1965b:246 [New Guinea]. Phatnoma vernoniae.—Drake and Ruhoff, 1965a:39. [NOTE.—The species Phatnoma amazonicum Drake and Hambleton [emendation]. name, derived from the generic name of the host plant, was originally Phatnoma amazonica Drake and Hambleton, 1944:120 [Brazil].—Drake and misspelled because it was based on the misspelled plant genus name given Ruhoff, 1965a:35. (as "Veronia") with the original description of this insect] Phatnoma annulipes Champion, 1897:4 [Panama].—Drake and Ruhoff, 1965a:35. Phatnoma annulipes annulipes Champion.—Drake, 1948b:21. Genus Phatnomella Pericart Phatnoma annulipes var. concision Drake [emendation]. Phatnoma annulata [sic] var. concisa Drake, 1948b:21 [Venezuela]. FIGURE 31 Phatnoma annulipes var. concisa.—Drake and Ruhoff, 1965a:35. Phatnomella Pericart, 1981:598 [type species: Phatnomella cristata Pericart, Phatnoma barberi Drake, 1941:141 [Colombia].—Drake and Ruhoff, 1965a: monobasic]. 36. Phatnoma biordinatum Froeschner, 1976:183 [Galapagos Islands]. DIAGNOSIS.—Among the genera with 8-9 head spines or Phatnoma costalis Distant, 1909:113 [Burma].—Drake and Ruhoff, 1965a:36. tubercles (occipitals much shorter than eye) this one can be Phatnoma coyazana Drake, 1948a:15 [Brazil].—Drake and Ruhoff, 1965a:36. recognized by the outer margin of the paranotum forming a Phatnoma ecuadore Drake [emendation]. Phatnoma ecuadoris Drake, 1941:141 [Ecuador].—Drake and Ruhoff, distinct, slightly acute angle anteriorly, and one opposite 1965a:36. humerus (this one with only 2 rows of cells) plus the 3-carinate Phatnoma eremaeum Drake and Froeschner [emendation]. pronotal disc. Length 1.8 mm. Phatnoma eremaea Drake and Froeschner, 1967:83 [Galapagos Islands]. DISTRIBUTION.—India. Phatnoma guatemalana Drake, 1948b:20 [Guatemala].—Drake and Ruhoff, ETYMOLOGY (feminine).—Phatnom-, a fragment of the 1965a:36. Phatnoma hacked Drake, 1950:154 [Australia].—Drake and Ruhoff, 1965a:36. generic name Phatnoma Fieber, plus ella, Latin (diminutive), Phatnoma hova Schouteden, 1957:82 [Madagascar].—Drake and Ruhoff, indicating a "small Phatnoma." 1965a:37. List of Phatnomella Species Phatnoma jinjana Drake, 1956:13 [Uganda].—Drake and Ruhoff, 1965a:37. Phatnoma laciniatum Fieber, 1844:57 [emendation; "Ostindien"].—Drake and Phatnomella cristata Pericart. 1981:599 [India]. Ruhoff, 1965a:37. Phatnomella variabilis Pericart, 1991:35 [Thailand]. 32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Phatnomella Species

Costal area on apical half 5-seriate. Dorsal margin of median pronotal carina equally high anterior and posterior to a deep rectangular or acute indentation near midlength P. variabUis Costal area (except at base) 3-seriate. Dorsal margin of median pronotal carina shallowly concave, anterior part distinctly higher than posterior part P. cristata

FIGURE 32.—Plesionoma humeralis, natural length 3.5 mm. FIGURE 31.—Phatnomella cristata, natural length 1.8 mm.

wide) into a prominent lobe. Length 3.1-3.5 mm. Genus Plesionoma Drake GEOGRAPHIC DISTRIBUTION.—This genus has been reported

FIGURE 32 only from Congo and South Africa in the southern half of Africa. Plesionoma Drake, 1950:157, 166 [type species: Phatnoma humeralis Distant, ETYMOLOGY (neuter).—plesios, Greek (near), plus fragment monobasic].—Drake and Runoff, 1965a:39. -noma from neuter generic name Phatnoma Fieber, suggesting DIAGNOSIS.—Among the genera with 8-9 head spines a relationship with Phatnoma. and/or tubercles this one can be recognized by the posterior COMMENTS.—Examination of the type of all species two-thirds of the paranotum being broadly expanded (5-6 cells cataloged in this genus by Drake and Ruhoff (1965a:39-40) NUMBER 574 33

revealed that eteosa Drake was misassigned; it is herein Plesionoma capeneri Duarte-Rodrigues, 1981b:2O3 [South Africa]. transferred to the genus Ulmus Distant. Plesionoma humerale (Distant) [emendation]. Phatnoma humeralis Distant, 1902a:239 [South Africa]. List of Plesionoma Species Plesionoma humeralis.—Drake and Runoff, 1965a:40. Plesionoma biseriatum Duarte-Rodrigues [emendation]. Plesionoma leroyi Schouteden, 1955a:25 [Congo].—Drake and Runoff, Plesionoma biseriatus Duarte-Rodrigues, 1987a: 176 [South Africa]. 1965a:40.

Key to Plesionoma Species

1. Head armature consisting of sharp spines as long as or longer than horizontal diameter of an eye. Lateral margin of paranotal dilation with 1 or more acute, prolonged spines 2 Head armature reduced to blunt tubercles shorter than horizontal diameter of an eye. Lateral margin of paranotum without spines 3 2. Costal area 3-seriate. Lateral projection of paranotum with 3 acute angulations P. humeralis Costal area 2-seriate. Lateral projection of paranotum with 1 acute angulation and 1 rounded lobe P. biseriatum 3. Costal area 6-seriate to apex of clavus, 4-seriate beyond P. leroyi Costal area 4-seriate to apex of clavus, 3-seriate beyond P. capeneri

Genus Pseudacalypta Pericart

FIGURE 33

Pseudacalypta Pericart, 1983:595 [type species: Pseudacalypta nepalensis Pericart, only included species]. DIAGNOSIS.—The arrangement of head spines or tubercles (presence of a dorsomedial and jugals and absence of occipitals and frontals) coupled with the strongly triangular bucculae (5-6 cells posteriorly, tapering to 1 cell anteriorly) mark this genus within the Phatnomatini. Length 2.4-2.5 mm. GEOGRAPHIC DISTRIBUTION.—The only species was re- ported from Nepal. ETYMOLOGY (feminine).—pseudo, Greek (false), plus Aca- lypta, generic name of small lace bugs in subfamily Tinginae that the member of the present genus resembles. List of Pseudacalypta Species Pseudacalypta nepalensis Pericart, 1983:5% [Nepal].

Genus PuUocader Pericart

FIGURE 34

PuUocader Pericart, 1991:38 [type species: Pullocader borneensis Pericart, monobasic]. DIAGNOSIS.—Among the genera with neither dorsomedial nor occipital armature, this genus will be recognized by bearing five tubercles (jugals, frontals, and a clypeal) on head, costal area widening from base, and 3-seriate pronotum. Length 1.7 mm. GEOGRAPHIC DISTRIBUTION.—Sabah. FIGURE 33.—Pseudacalypta nepalensis, natural length 2.4 mm. ETYMOLOGY (masculine).—pullus, Latin (dark colored), 34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

recognized by having the paranotum somewhat oblique (not recurved above itself) and with outer margin simple (no spines or angular projections, and the peritreme obscure, not forming a distinctly elevated, apically closed loop. Length 2.5-3.9 mm. GEOGRAPHIC AND GEOLOGIC DISTRIBUTION.—The modern species of this genus are confined to southern Africa, where their range extends from Kenya south to the southern tip of the continent. A fossil specimen imbedded in amber designated as "Baltic" in the original description (but without supporting evidence) may indicate a wider geographic range earlier or may be mislabeled as to origin. ETYMOLOGY (feminine).—G.E. Steyskal interpreted this term as a newly coined word. COMMENTS.—Two species not appearing in the Drake and Ruhoff list for Gonycentrum are added to this genus: a recently described species, G. haplotaxis Froeschner, and the present transfer of the fossil species baltica Drake from Phatnoma; the latter move was made necessary when examination revealed that the head of the type specimen in amber clearly shows a dorsomedial tubercle, a structure not present in species of the genus Phatnoma, the presence of occipitals, three carinae on pronotal disk, and the 1- or 2-seriate paranotum.

List of Sinalda Species Sinalda aethiops (Distant).—Froeschner, 1968:249. [Note: Distant's (1904b:427) assignment of this species to the genus Sinalda was supported by Froeschner's (1968, above) studies.] Phatnoma aethiops Distant, 1902:238 [South Africa].

FIGURE 34.—Pullocader borneensis, natural length 1.7 mm. plus nondescriptive -cader, fragment of generic name Canta- cader to indicate another genus in the taxon containing that genus, a comment on a Ca/i/aca

List of Pullocader Species Pullocader borneensis Pericart, 1991:38 [Sabah].

Genus Sinalda Distant

FIGURE 35

Sinalda Distant, 1904b:426 [type species: Sinalda elegans Distant, subsequent designation by Monte, 1947:4]. [Note: This taxon was cataloged by Drake and Runoff (1965a:32) as a junior synonym of Gonycentrum but was returned to genus status by Froeschner (1968:248). See "Discussion" under Gonycentrum.]

DIAGNOSIS.—Among those genera of the tribe Phatnomini bearing 8-9 spines or tubercles on the head, Sinalda may be FIGURE 35.—Sinalda elegans, natural length 2.5 mm. NUMBER 574 35

Gonycentrum aethiops.—Drake and Ruhoff, 1965a:32. Sinalda afra (Drake and Ruhoff).—Froeschner 1968:249. Gonycentrum afrum Drake and Ruhoff. 1961:126 [South Africa]; 1965a: 31. Sinalda angustata (Drake).—Froeschner, 1968:249. Gonycentrum angustatum Drake, 1956:15 [Tanzania].—Drake and Ruhoff, 1965a:32. Sinalda baltica (Drake) [fossil! new combination]. Phatnoma baltica Drake, 1950:153 [Baltic amber].—Drake and Ruhoff, 1965a:35. Sinalda capensis Duarte-Rodrigues, 1988:494 [South Africa]. Sinalda cristata Duarte-Rodrigues, 1988:495 [South Africa]. Sinalda dilatata Duarte-Rodrigues, 1988:496 [South Africa]. Sinalda elegans Distant, 1904b:427 [South Africa].—Froeschner 1968:249. Gonycentrum elegans.—Drake and Ruhoff, 1965a:33. Sinalda haplotaxis Froeschner, 1968:249 [Transvaal]. Sinalda helichrysumae Duarte-Rodrigues, 1981b:206 [South Africa]. Sinalda nebulosa Distant, 1904b:428 [South Africa].—Froeschner. 1968:249. Gonycentrum nebulosum.—Drake and Ruhoff, 1965a:33. Sinalda reticulata Distant, 1904b:427 [South Africa].—Froeschner, 1968:249. Gonycentrum reticulatum.—Drake and Ruhoff, 1965a:33. Sinalda sinuaticollis (Linnavuori).—Duarte-Rodrigues, 1981:207. Gonycentrum sinuaticolle Linnavuori, 1977:6 [Ethiopia]. Sinalda testacea (Distant).—Froeschner, 1968:249. Phatnoma testacea Distant, 1902a:238 [South Africa]. Gonycentrum testaceum.—Drake and Ruhoff, 1965a:34. Sinalda thomasi (Drake).—Froeschner, 1968:249. Gonycentrum thomasi Drake, 1956:14 [Kenya].—Drake and Ruhoff, 1965a:34.

Genus Taphnoma Pericart FIGURE 36

Taphnoma Pericart, 1991:42 [type species: Taphnoma brunneicornis Pericart, original designation]. DIAGNOSIS.—Among those genera of Phatnomatini with jugals, frontals, dorsomedial, and one on clypeus, but no evident occipitals, Taphnoma is recognized by the 1-carinate pronotal disc plus the free margin of the paranotum simple or forming a prominent angulation opposite humerus. Length 2.3-3.1 mm. GEOGRAPHIC DISTRIBUTION.—Known only from southern China and the island of Borneo. ETYMOLOGY (neuter).—This name is an anagram of the generic name Phatnoma. FIGURE 36.—Taphnoma brunneicornis, natural length 3.1 mm. COMMENT.—The present assignment of Malala tuberculum Jing to this genus resulted from personal examination of the holotype.

Lists of Taphnoma Species Taphnoma elegans Pericart, 1991:46 [Sabah]. Taphnoma tuberculum (Jing) [new combination]. Taphnoma acutispinis Pericart, 1991:44 [Sabah]. Malala tuberculum Jing, 1980:400,403 [China]. Taphnoma brunneicornis Pericart 1991:43 [Sabah].

Key to Taphnoma Species

Costal area 3-seriate on basal two-thirds or more. Paranotal margin distinctly angled opposite humerus 2 Costal area mostly 2-seriate, 3-seriate only in basal fifth. Pronotal margin broadly, convexly rounded opposite humerus 3 36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

2. Head black, pronotum (except calli) yellow brown. Discoidal and subcostal areas with 2-3 darkened elevated cross veins T. brunneicornis Head and anterior thirds of pronotum and paranota pale; posterior lobe of pronotum and posterior two-thirds of paranotum black T. acutispinis 3. Paranotum narrower than an eye; its free margin virtually straight except for curving inward opposite humerus T. tuberculum Paranotum distinctly wider than an eye; its free margin broadly convex for full length T. elegans

Genus Thaicader Pericart

FIGURE 37

Thaicader Pericart, 1991:40 [type species: Thaicader burckhardti Pericart, monobasic]. DIAGNOSIS.—The presence of a single carina on the pronotal disc and a dorsomedial spine, coupled with the absence of jugals, combine to separate this genus from all other Phatnomatini genera except Pseudacalypta Pericart; the latter genus has a broadly triangular buccula (4-5 cells wide posteriorly, tapering to a single cell anteriorly) while in Thaicader the buccula is 2-seriate for nearly full length. Length 2.3-2.6 mm. GEOGRAPHIC DISTRIBUTION.—This genus is known only from Thailand. ETYMOLOGY (masculine).—Thai-, from Thailand, country of origin, plus -cader, nondescriptive, from genus Cantacader to indicate yet another genus in the taxon containing that genus. List of Thaicader Species Thaicader burckhardti Pericart, 1991:40 [Thailand].

Genus Ulmus Distant

FIGURE 38

Ulmus Distant, 1904b:426 [type species: Ulmus testudineatus Distant, monobasic].—Drake and Runoff, 1965a:41. DIAGNOSIS.—Among the genera of the tribe Phatnomini, with 8-9 cephalic spines or tubercles (each occipital at least as long as an eye), Ulmus may be recognized by the presence of a spine on midline of head between the frontals (anterior to dorsomedial). Length 2.2-3.7 mm. GEOGRAPHIC DISTRIBUTION.—This genus has been reported from Africa south of the Sahara Desert. ETYMOLOGY (feminine).—ulmus, Latin (elm), of unknown significance as applied to these insects. COMMENTS.—Redefinition of genera brought about several changes in the cataloged contents of this genus. Of the two species listed by Drake and Ruhoff (1965a:41) only the type species testudineatus Distant remains; Drake and Ruhoff s U. engaeus is herein removed to become the type species of the new genus Exulmus. In addition, two species, Plesionoma FIGURE 37.—Thaicader hurckhardti, natural length 2.3 mm. eteosa Drake and P. drakei Schouteden, are herein added by NUMBER 574 37 transfer from the genus Plesionoma. Ulmus eteosa (Drake) [new combination]. Plesionoma eteosa Drake, 1954:2 [South Africa].—Drake and Runoff, List of Ulmus Species 1965*39. Ulmus drakei (Schouteden) [new combination]. Ulmus testudineatus Distant, !904b:426 [Transvaal].—Drake and Runoff, Plesionoma drakei Schouteden, 1965b:353 [Kenya]. 1965*41.

Key to Ulmus Species

1. Margin of paranotum with an acute, spine-like projection anterolaterally 2 Margin of paranotum obtusely angled (no prolonged spine)....(/. testudineatus 2. Clypeus with two long, erect spines (midlength and subapically). Costal area 4-seriate on basal third, 3-seriate beyond U. eteosa Clypeus with 1 erect spine (midlength). Costal area mostly 3-seriate, 2-seriate at midlength and apex U. drakei

Genus Zetekella Drake

FIGURE 39

Zetekella Drake, 1944:139, 142 [type species: Zetekella zeteki Drake, monobasic].—Drake and Ruhoff, 1965a:41.

DIAGNOSIS.—Among those genera of the tribe Phatnomatini lacking the dorsomedial spine or tubercle, Zetekella can be recognized by its ostiolar pore being so close to the hypocosta that its rim overlaps the ventral vein of the hypocosta (no extended peritreme present). Length 1.8-2.0 mm. GEOGRAPHIC DISTRIBUTION.—The known species occur only in tropical America. ETYMOLOGY (feminine).—A patronym for James Zetek, collector of the type material, plus -ella, Latin (diminutive). COMMENTS.—The present treatment differs from the Drake and Ruhoff Catalog (1965a:41) by considering Zetekella and Minitingis as separate and valid genera, the latter being a West Indies genus, the former confined to continental tropical America. Additional discussion is presented under the treat- ment of Minitingis. The present illustration of the type species was made from the sketch accompanying the original description of Z. zeteki; the holotype of that species was subsequently badly damaged (now lacks head and both paranota) by dermestids. Dr. Karol Leno, Sao Paulo, Brazil, in a personal note, reported the observation that Zetekella pulla Drake and Hambleton is "very common in the nests of ants Camponotus rufipes and Odontomachus affinis as well as in the forest humus."

List of Species of Zetekella Zetekella pulla Drake and Plaumann, 1956:17 [Brazil].—Drake and Ruhoff, 1965a:41. Zetekella zeteki Drake, 1944:140 [Panama].—Drake and Ruhoff, 1965a:41. FIGURE 38.—Ulmus testudineatus, natural length 2.7 mm. 38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Zetekella Species

Paranota very wide, width subequal to width of head, with 4 to 5 rows of cells; costal area with 4 rows of cells for full length Z. zeteki Paranotum much narrower, about half as wide as head, with only 2 rows and a few intercalary cells; costal area with only 2 rows of cells Z. pulla

FIGURE 39.—Zetekella zeteki, natural length 2.0 mm. Literature Cited

Amyot, CJ.B., and J.G.A. Serville 1941. New American Tingitidae (Hemiptera). Journal of the Washington 1843. Histoire naturelle des insectes: Himiptires. lxxvi + 675 + 8 pages, Academy of Sciences, 31:141-145. 12 plates. Paris: Fain et Thunot. 1944. Concerning the American Cantacaderinids (Hemiptera: Tingidae). Barber, H.G. Boletin de Entomologia Venezolana, 3:139-142. 1954. A Report on the Hemiptera-Heteroptera from the Bimini Islands, 1947. Tingidae (Hemiptera) from the Orient and South Pacific. Musee Bahamas, British West Indies. American Museum Novitates, Heude, Notes d'Entomologie Chinoise. 11:225-231. 1682:1-18. 1948a. Five New American Tingidae (Hemiptera). Boletin de Entomologia Bekker-Migdisova, E.E. Venezolana, 7:15-19. 1962. [Order Heteroptera; True Bugs.] In B.B. Rohdendorf, editor, 1948b. New American Tingidae (Hemiptera). Boletin de Entomologia Fundamentals of Paleontology, 9:289-317. [In Russian; translated Venezolana. 7:20-25. into English, 1991, edited by D.R. Davis. Translations Publishing 1950. Concer ning the Can tacaderi nae of the World (Hemiptera: Tingidae). Program, Smithsonian Institution Libraries, Natural History Build- Arthropoda, 1:153-166. ing, Washington, D.C. 20560]. 1951. New Genera and Species of Tingidae (Hemiptera) in the Hungarian Bergroth, E. National Museum. Annales Historico-Naturales Musei Nationalis 1894. Tingidae tres Madagascariensis. Revue d'Entomologie, 13:167- Hungarici, 1:165-178. 168. 1954. Tingidae: Descriptions and Synonymic Data (Hemiptera). Great 1898. Eine neue Tingide. Wiener Entomologische Zeitung, 17:9. Basin Naturalist, 14:1-10. 1908. Neue Hemiptera aus Sud-Abyssinien. Revue Russe d'Entomologie, 1956. Three New Species of Cantacaderinae from Africa (Hemiptera: 1907:106-110. Tingidae). Revue de Zoologie et de Botanique Africaines, 53: 1927 ("1926"). Hemiptera Heteroptera from New Zealand. Transactions 13-16. and Proceedings of the New Zealand Institute, 57:671-684. 1957. Quelques Tingidae de la Reunion (Hemiptera). Memoirs de B16te, H.C. I Institute Scientifique de Madagascar, 8:399-405. 1945. Catalogue of the Berytidae, Piesmidae and Tingidae in the 1960. Tingidae of New Guinea (Hemiptera). Pacific Insects. 2:339-380. Rijksmuseum van Natuurlijke Historic Zoologische Mededeelingen, 1961. A New Genus and Species of Cantacaderine Lace-bug from the 25:72-92. Philippines (Hemiptera: Tingidae). Fieldiana (Zoology), 42:115- Bruner, S.C. 118. 1940. A New Tingitid from Cuba (Hemiptera). Memorias de la Sociedad Drake, C.J., and N.T. Davis Cuba de Historia Natural, 14:245-247. 1960. The Morphology, Phytogeny, and Higher Classification of the Champion, G.C. Family Tingidae, Including the Description of a New Genus and 1897. Tingitidae. In Godman and Salvin, editors, Biologia Centrali- Species of the Subfamily Vianaidinae (Hemiptera: Heteroptera). Americana, 2:1-48. Entomologica Americana, 39:1-100. Distant, W.L. Drake, C.J., and R.C. Froeschner 1902a. Rhynchotal Miscellanea. Annals of the South African Museum, 1967. Lacebugs of the Galapagos Archipelago (Hemiptera: Tingidae). 2:237-254, plate XV. Proceedings of the Entomological Society of Washington. 69:82-93. 1902b. Rhynchotal Notes, XIII; Heteroptera: Families Tingitidae, Phymati- Drake, C.J., and E.J. Hambleton dae, and Aradidae. Annals and Magazine of Natural History, series 1934. Brazilian Tingitidae (Hemiptera), Part I. Revista de Entomologia. 7, 9(59):353-362. Rio de Janeiro, 4:435-451. 1903-1904a. Rhynchota. In William T. Blanford, editor. The Fauna of 1938. Brazilian Tingitoidea (Hemiptera), Part IV. Revista de Entomologia, British India, Including Ceylon and Burma. Volume 2: 1-242 Rio de Janeiro, 5:51-57. (1903); 143-503 (1904). London: Taylor and Francis. 1944. Concerning Neotropical Tingitidae (Hemiptera). Journal of the 1904b. On the South African Tingididae and Other Heteropterous Rhyn- Washington Academy of Sciences, 34:120-129. chota. Transactions of the South African Philosophical Society, Drake, C.J., and T. Maa 14:425-436,1 plate. 1955. Chinese and Other Oriental Tingitoidea (Hemiptera), III. Quarterly 1909. New Oriental Tingididae. Annales de la Sociiti Entomologique de Journal of the Taiwan Museum, 8:1-11. Belgique, 53:113-123. Drake, C.J., and F. Plaumann 1910. Rhynchota (Heteroptera: Appendix). In William T. Blanford, editor. 1956. A New Cantacaderid from Brazil (Hemiptera: Tingidae). Bulletin of The Fauna of British India, Including Ceylon and Burma. Volume 5: the Southern California Academy of Sciences, 55:14-18. 100-126. London: Taylor and Francis. Drake, C.J., and M.E. Poor Drake, C.J. 1936. New Indian Tingitidae (Hemiptera). Indian Forest Records, 2:141- 1922. Neotropical Tingitidae with Descriptions of Three New Genera and 145. Thirty-two New Species and Varieties (Hemiptera). Memoirs of the Drake, C.J.. and F.A. Runoff Carnegie Museum, 9:351-377, plate 39. 1961. New Genera and New Species of Lacebugs from the Eastern 1923. Two New Species of Cantacaderia (Hemip. Tingitidae). Bulletin of Hemisphere (Hemiptera: Tingidae). Proceedings of the United the Brooklyn Entomological Society, 18:81-84. States National Museum, 113:125-183. 1928. New and Little Known Neotropical Tingitidae. Iowa State College 1962. Some Tingidae (Hemiptera) in the South Australian Museum. Journal of Science, 3:41-56. Records of the South Australian Museum, 14:249-252.

39 40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1965a. Lacebugs of the World, a Catalog (Hemiptera: Tingidae). United 1943b. Form of Family Name to be Formed from the Name Tingis Fabricius States National Museum Bulletin, 243:i-viii, 1-634, frontispiece, (Hemiptera). Bulletin of Zoological Nomenclature, 1:13. plates 1-56. Jing, X.-L. 1965b. Lacebugs from New Guinea, Borneo, Solomons, and Other Islands 1980. [New Species of Chinese Tingidae (Hemiptera: Heteroptera).] Acta of the South Pacific and Indian Oceans (Hemiptera: Tingidae). Zootaxonomica Sinica, 5:395-403. [In Chinese, with English Pacific Insects, 7:243-290. summary on pages 402-403.] Duarte-Rodrigues, P. Kirkaldy, G.W. 1978. African Tingidae, II: Descriptions of Three New Species (Heterop- 1908. A Catalogue of the Hemiptera of Fiji. Proceedings of the Linnean tera). Boletim Sociedade Portuguesa de Ciencias Naturais, 18:13- Society of New South Wales, 33:345-391. 18. Lepeletier, A.L.M., and J.G.A. Serville 1980. African Tingidae, XIII: A New Species of Cantacader Amyot and 1825-1828. [Articles on Hemiptera.] In G.A. Olivier, editor, Encyclopidie Serville from Malawi (Heteroptera). Arnoldia Zimbabwe, 80(40): 1- Mithodique, volume 10, 833 pages. Paris: Agasse. [Pages 1-344 3. published in 1825; 345-833 in 1828.] 1981a African Tingidae, XXI: Lacebugs in the British Museum (Natural Linnavuori, R. History) (Heteroptera). Arquivos do Museu Bocage, series C, 1977. Hemiptera of the Sudan, with Remarks on Some Species of the 1:133-200. Adjacent Countries, 5: Tingidae, Piesmidae, Cydnidae, Thaumastel- 1981b. African Tingidae, XXII: Lacebugs in the Plant Protection Institute lidae and Plataspidae. Acta Zoologica Fennica, 147:1-81. (Pretoria) (Heteroptera). Arquivos do Museu Bocage, series C, Montandon, A.L. 1:201-256. 1892. He'mipteres-He'te'ropteres nouveaux. Revue d'Entomologie, 11: 1982. African Tingidae, XXV: A New Cantacader, Three New Agramma 265-273. and New Data (Heteroptera). Arquivos do Museu Bocage, series A, Monte, O. 1:325-334. 1942. Critica sobre alguns generos e espe'cies de Tingitideos. Papiis 1987a. African Tingidae, XVIII: Three New Species and New Data from Avulsos do Departmento de Zoologia, Sao Paulo, 2:103-115. South and Southwest Africa (Heteroptera). Arquivos do Museu 1946. Sobre o genero Phatnoma com a descricao de uma nova esp6cie e a Bocage, series B, 2(21): 175-185. lista da suas espe'cies (Hem. Tingidae). Revista Brasileira de 1987b. New Species and Records of Lacebugs (Heteroptera: Tingidae) from Biologia, 6:247-254. Southern Africa. Annals of the Transvaal Museum, 34:349-369. 1947. Generos e genotipos dos Tingideos do mundo. Papiis Avulsos do 1988. African Tingidae, XXVIII: New Species and New Data from South Departmento de Zoologia, Sao Paulo, 8:1-22. and South West Africa. Boletim da Sociedade Portuguesa de Myers, J.G. Entomologia, 67:493-515 (1984). 1922. The Order Hemiptera in New Zealand, with Special Reference to Its Fieber. F.X. Biological and Economic Aspects. The New Zealand Journal of 1844. Entomologische Monographien. 138 pages, 10 plates. Leipzig: J.A. Science and Technology, 5:1-12. Barth. 1926. Biological Notes on New Zealand Heteroptera. Transactions of the Froeschner, R.C. New Zealand Institute, 56:449-511. 1968. Notes on the Systematics and Morphology of the Lacebug Subfamily Myers. J.G., and W.E. China Cantacaderinae. Proceedings of the Entomological Society of 1928. A List of New Zealand Heteroptera with Description of a Washington, 70:245-254. Remarkable Green Aradid Representing a New Genus. Annals and 1976. Galapagos Lace Bugs: Zoogeographic Notes and a New Species of Magazine of Natural History, series 10, 1:377-394. Phatnoma (Hemiptera: Tingidae). Proceedings of the Entomological Pericart, J. Society of Washington. 78:181-184. 1981. Quatre especes nouvelles de Cantacaderinae du Nord de l'lnde, 1981. Heteroptera or True Bugs of Ecuador A Partial Catalog. Smith- repre'sentant trois genres nouveaux (Hemiptera: Tingidae). Revue sonian Contributions to Zoology, 322:1-147. Suisse de Zoologie, 88:595-605. Germar, E.F., and G.C. Berendt 1983. Deux Cantacaderinae nouveaux du Ne'pal de la tribu des Phatnomini 1856. Die im Bernstein befindlichen Hemiptera und Orthopteren der (Hemiptera: Tingidae). Revue Suisse de Zoologie, 90:593-597. Vorwelt. In G.C. Berendt, Die im Bernstein befindlichen organis- 1986. Froeschnerocader denticollis (Hetroptera: Tingidae): A New Genus chen Reste der Vorweld. 2:1-40, plates 1-4. and Species of Cantacaderinae from Bomo. Journal of the New York Hacker. H. Entomological Society, 94:245-248. 1927. New Tingitoidea (Hemiptera) in the Queensland Museum. Memoirs 1991. Cantacaderinae de Thailande, Borneo et Palawan: Genres nouveaux, of the Queensland Museum, 9:19-22, plates 6-10. especes nouvelles ou inte'ressantes ainsi qu'une nouvelle synonymie 1928. New Species and Records of Australian Tingitoidea (Hemiptera). (Hemiptera, Tingidae). Revue Suisse de Zoologie, 98:33-50. Memoirs of the Queensland Museum, 9:174-188, plates 20-23. Schouteden, H. 1929. New Species of Australian Tingitidae (Hemiptera). Memoirs of the 1916. Tingides du Congo Beige. Revue de Zoologie Africaine, 4:288-297. Queensland Museum, 9:324-334, plates 32-35. 1923. Nouvelles notes sur les Tingidae du Congo Beige. Revue de Zoologie Henning, W. Africaine. 11:82-110. 1966. Phylogenetic Systematics. ii + 263 pages. Urbana, Illinois: Univer- 1955a. Tingides nouveaux du Congo Beige. Revue de Zoologie et de sity of Illinois Press. Botanique Africaines, 52:25-32. Horv&th,G. 1955b. Tingides nouveau des collections du Musee Royal du Congo Beige. 1906. Synopsis Tingitidarum Regionis Palaearcticae. Annales Musei Revue de Zoologie et de Botanique Africaines, 52:162-168. Nationalis Hungarici, 4:1-118. 1957. Tingides de Madagascar. Revue de Zoologie et de Botanique International Commission of Zoological Nomenclature Africaines, 55:82-89. 1943a. Opinion 143: On the Method of Forming the Family Name for Tingis 1965a. Tingides Africaines nouveaux de genre Cantacader Amyot et Fabricius, 1803 (Insecta. Hemiptera). Opinions and Declarations Serville. Revue de Zoologie et de Botanique Africaines, 72:168-172. Rendered by the International Commission on Zoological Nomen- 1956b. Plesionoma Drakei nov. spec. Tingide nouveau d'Afrique orientate clature. 2:83-87. (Hem.). Revue de Zoologie et de Botanique Africaines, 72:353-354. NUMBER 574 41

Scott, J. 11(2):1-163; part 4 (1874). 12(1):1-186; part 5 (1876), 14(4):1- 1874. On a Collection of Hemiptera Heteroptera from Japan; Descriptions 162. of Various New Genera and Species. Annals and Magazine of Stusak, J.M. Natural History, series 4, 14:289-304, 360-365,426-452. 1976. A New Genus and Species of Cantacaderinae from Java (Heterop- Scudder, G.G.E. tera, Tingidae). Ada Entomologica Bohemoslovaca, 73:13-16, plate 1959. The Female Genitalia of the Heteroptera: Morphology and Bearing L on Classification. Proceedings of the Royal Entomological Society of 1979. Cantacaderinae Collected by the Hungarian Expedition to West London, 111:405-467. Afrika with Some Notes on Cantacaderinae (Heteroptera, Tingidae). Scudder, S.H. Opuscula Zoo logic a Budapest, 16:141-149. 1890. The Tertiary Insects of North America. United States Geological 1984. Two New Species of Afrotropical Tingidae (Heteroptera). Ada Survey Bulletin, 71:1-663. Faunistica Entomologica Musei Nationalis, Pragae, 17:237-243. Spinola, M. Stys, P., and I.M. Kerzhner 1852. Hemipteros. In C. Gay, editor, Historia fisica y politica de Chile. 1975. The Rank and Nomenclature of Higher Taxa in Recent Heteroptera. Zoo/og/a. 7:113-320. Ada Entomologica Bohemoslavaca, 72(2):65-79. Stal.C. Takeya, C. 1865-1866. Hemiptera Africana. Volumes 1-4. Holmie: Officina Nor- 1933. New or Little-known Lace Bugs from Japan, Korea and Formosa stedtiana [1865, i:iv + 256; 1866, 2:1-181; 3:1-200; 4:i + 275 + I]. (Hemiptera: Tingitidae). Mushi, 6:32-39. 1870-1876. Enumeratio Hemipterorum: Bidrag till en Foreteckning ofver Woodward, T. alia hittils kanda Hemiptera, jemte systematiscka meddelanden, 1961. The Heteroptera of New Zealand, Part HI: Coreidae, Berytidae, Parts 1 -5. Kongliga Svenska Vetenskaps-Akademiens Handlingar, Tingidae, Cimicidae. Transactions of the Royal Entomological part 1 (1871), 9(l):l-232; part 2 (1872), 10(4):l-159; part 3 (1873), Society of New Zealand, 1:145-158. Index

(Synonyms and page numbers of principal accounts in italics) abdivitus, Cantacader, 8 Cantacaderini, 3, 4, 5. 10, 11, 17, 23, 31 Ulmus, 36, 37 Acalypta,3 capeneri, Plesionoma, 33 Etesinalda, 5, 17, 18,24 acalytoides, Cyclotynaspis, 21,22 capensis, Sinalda, 35 Exulmus, 5, 17, 18,25 acutispinis, Taphnoma, 35, 36 capitatus, Astolophus, 20 formosus, Cantacader, 9 aethiops, Gonycentrum, 35 carectorum, Cyperobia, 10-12 froeschneri, Phatnocader, 29, 30 Phatnoma, 34 Carldrakeana, 5, 6, 9, 10, 26 Froeschnerocader, 15, 16 Sinalda, 34 Ceratocader, 5, 6, 10 furtivus, Cantacader, 9 afra, Sinalda, 35 chadense, Gonycentrum, 26 charieis, Distocader, 22, 23 afirum, Gonycentrum. 35 gerardi, Cantacader, 9 afzelii, Cantacader, 8 Malala, 22, 23 Gonycentrum. 5.9, 17, 19, 23, 25, 26, 34 agilis, Cantacader, 8 clairi, Cantacader, 8 gounellei, Nectocader, 12, 13 agviates, Phatnoma, 29, 31 claratis, Cantacader, 8 Cantacader. 12, 13 ainata, Phatnoma, 31 Cnemiandrus, 5, 19, 21 guatemalana, Phatnoma, 31 ainatum, Phatnoma, 31 concisa, Phatnoma, 31 allaeri, Cantacader, 8 concisum, Phatnoma, 31 hackeri, Phatnoma, 31 Allocader, 5, 6, 7 cordata, Phatnoma, 7 haplotaxis, Sinalda, 34, 35 Alloeoderes, 5. 18,19 cordatus, Allocader, 7 helichrysumae, Sinalda, 35 amazonica, Phatnoma, 31 comiculata, Pseudophatnoma, 15, 16 Hcteroptera, I amazonicum, Phatnoma, 31 coronata, Teleia, 25, 26 nova, Phatnoma, 31 amplicostatus, Cantacader, 8 coronatum, Gonycentrum, 26 hulstaerti, Cantacader, 9 amydis, Cantacader, 8 costalis, Phatnoma, 31 humerale, Plesionoma, 33 Angiocader, 5, 17, 18,20 coyazana, Phatnoma, 31 humeralis, Phatnoma, 32, 33 angulipennis, Cantacader, 8 cristata, Phatnomella, 31, 32 humeralis, Plesionoma, 33 angustata, Sinalda, 35 Sinalda, 34, 35 angustalum, Gonycentrum, 35 curtulus, Cantacader, 8 ilongaensis, Cantacader, 9 angustecostatus, Cantacader, 8 Cyclotynaspis, 5, 19, 21 Indocader, 5, 17, 18,26 annulipes, Phatnoma, 31 Cyperobia, 5, 6, 11, 17 infuscatus, Cantacader, 9 antennata, Eotingis, 14 insularis, Cantacader, 9 armatus, Cantacader. 10 Daillea.5, 17, 18,22 japanicus, Cantacader, 9 Ceratocader, 10, 11 davao, Alloeoderes, 19 jinjana, Phatnoma, 31 Astolophus, 5, 19,20 dentatus, Ceratocader, 10 attenuatus, Cantacader, 8 Cantacader, 10 laciniata, Phatnoma, 30, 31 avitus, Cantacader, 1,6, 14 denticollis, Froeschnerocader, 15, 16 laciniatum, Phatnoma, 31 Paleocader, 13,14 Pseudophatnoma, 16 laratanus, Cantacader, 9 diffidentis, Cantacader, 9 laticollis, Cantacader, 9 baltica, Phatnoma. 1, 31, 35 dilatata, Sinalda, 35 laticosta, Etesinalda, 17,24 Sinalda, 1, 35 Distocader, 5,17, 18,22,23 leai, Cantacader, 7 barberi, Phatnoma, 31 divisus, Cantacader, 9 Allocader, 7 basilewskyi, Cantacader, 8 drakei, Plesionoma, 36, 37 leroyi, Plesionoma, 33 besucheti, Indocader, 26, 27 Ulmus, 36, 37 letabanus, Cantacader, 9 biordinatum, Phatnoma, 31 lethierryi, Cantacader, 9 ecuadore, Phatnoma, 31 biroi, Oranoma, 29 loebli, Indocader, 26, 27 ecuadoris, Phatnoma, 31 biseriatum, Plesionema, 33 longicomis, Cantacader, 9 biseriatus, Plesionema, 33 elegans, Gonycentrum. 35 bomeensis, Pullocader, 33, 34 Sinalda, 34, 35 maculate, Phatnoma, 31 bouclei. Montea. 23 Taphnoma, 35, 36 maculatum, Phatnoma, 31 Eocader. 23 elsae, Minitingis, 29 magnificus. Cantacader, 16 bowmansi, Cantacader, 8 engaeus, Exulmus, 17,25 Teratocader, 13, 16, 17 bninneicomis, Taphnoma, 35, 36 Ulmus, 25, 36 Malala, 23, 25, 26 bulliens. Malala. 25, 26 engista, Carldrakeana, 10 marmorata, Phatnoma, 31 burckhardti, Thaicader, 36 engistum, Gonycentrum. 10 marmoratum, Phatnoma, 31 Eocader, 5, 17.2J maynei, Phatnoma, 31 Cantacader, 5,6,8, 10, 14, 16,20,26,27, Eotingis, 14 Microcader, 5, 18,27 30,31.34.36 eremaea, Phatnoma, 31 Minitingis, 5, 17,19, 28, 29, 37 Cantacaderaria, 5 eremaeum, Phatnoma, 31 minuscula, Zetekella, 29 Cantacaderinae. 1, 3,4, 5,6, 14, 20, 30 eteosus. Plesionoma, 31, 36, 37 minusculus, Minitingis, 28, 29

42 NUMBER 574 43

Montea, 23 Taphrostethus, 9 tingidoides, Piesma, 16 Stenocader, 16 nebulosa, Sinalda, 35 reticulata, Sinalda, 35 Tinginae, 1, 3,4, 20, 33 nebulosum, Gonycentrum, 35 reticulation, Gonycentrum, 35 Tingini, 8,9 Nectocader, 5, 6,12, 13 Tingis, 28 nepalensis, Pseudacalypta, 33 schoutedeni, Cantacader, 9 Tingitidae, 4 nesiotes, Allocader, 7 sejunctus, Cantacader, 9 togulare, Phatnoma, 31 nocturnis, Cantacader, 9 Sinalda, 5, 18,24,26,31,54 togularis, Phatnoma, 31 nubilus, Cantacader, 9 sinuaticollis, Gonycentrum, 26, 35 tonkinana, Phatnoma, 31 Sinalda, 26, 35 tricarinatus, Cantacader, 8 socia, Carldrakeana, 10, 12 obesa, Phatnoma, 20 tricostata, Dailiea, 22 socium, Gonycentrum, 10 obesus, Angiocader, 20 trinidadana, Phatnoma, 31 Stenocader, 5, 6, 8,16, 17 Oranoma, 5,18, 29 tuberculum, Malala, 26, 35 Taphnoma, 26, 35, 36 ovata, Phatnoma, 31 takasago, Phatnoma, 31 typicus, Cnemiandrus, 21 ovatum, Phatnoma, 31 Taphnoma, 5, 17, 18, 26, 35 Teleia, 25 uichancoi, Phatnoma, 31 pacifica, Phatnoma, 31 tener, Cantacader, 9 Ulmus, 5, 18, 33.56 Paleocader, 1, 6, 8, 13 tenuipes, Cantacader, 9 unicarinata, Phatnoma, 29, 31 Phatnocader, 5, 17, 18, 29 Teratocader, 5, 6, 13,16 unicarinatum, Phatnoma, 31 Phatnoma, 5, 17, 18, 24,30, 31, 35 testacea, Phatnoma, 35 unicostatus, Microcader, 27,28 Phatnomatini, 4-6, 10, / 7. 23, 24, 26. 27, testacea, Sinalda, 35 uniformis, Cantacader, 9 30-32, 35-37 testaceum, Gonycentrum, 35 Phatnomella, 5, 18,5/ testudinatus, Ulmus, 36, 37 vandenplasi, Cantacader, 9 Phatnomini, 11, 18 thai, Microcader, 27, 28 variabilis, Phatnomella, 31, 32 Plesionoma, 5, 18,52,37 Thaicader,5, 17,18,56 varians, Phatnoma, 31 Pseudacalypta, 5. 17, 18,33, 36 thomasi, Gonycentrum, 25 variegatus, Microcader, 27,28 Pseudophatnoma, 5, 6,15, 17 Sinalda, 35 vergrandis, Eocader, 33 pulla, Zetekella, 37. 38 tindaiei, Carldrakeana, 9, 10 veridica, Phatnoma, 31 Pullocader, 5. 17, 19,33 Gonycentrum, 10 veridicum, Phatnoma, 31 Phatnoma, 9 vernoniae, Phatnona, 31 quadricornis, Piesma, 8, 9 Tingidae, 1,4, 14 veroniae, Phatnoma, 31 Cantacader, 3, 9 Tingideae, 4 Vianaididae, 1,2 quinquecarinata, Tingis, 6, 14 Tingidida, 4 Vianaidinae, 1 quinquecarinatus, Cantacader, 1, 8 Tingididae, 4 Paleocader, 13,14 Tingiditae, 4 Zetekella, 5, 19, 28,37 quinquecostatus, Cantacader, 9 Tingidites, 4 zeteki, Zetekella, 37, 38

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Illustrations that are intended to follow the printed text should be submitted as carefully typed, double-spaced copy may be termed Plates, and any components should be similarly separate from the text; they will be typeset unless otherwise lettered and referenced: "Plate 9b." Keys to any symbols within an requested. If camera-copy use is anticipated, do not draw rules on illustation should appear on the art rather than in the legend. manuscript copy. Some points of style: Do not use periods after such abbrevia- Taxonomic keys in natural history papers should use the tions as "mm, ft, USNM, NNE." Spell out numbers "one" through aligned-couplet form for zoology and may use the multi-level indent "nine" in expository text, but use digits in all other cases if possible. form for botany. If cross referencing is required between key and text, Use of the metric system of measurement is preferable; where use of do not include page references within the key, but number the the English system is unavoidable, supply metric equivalents in keyed-out taxa, using the same numbers with their corresponding parentheses. Use the decimal system for precise measurements and heads in the text. relationships, common fractions for approximations. Use day/month/ year sequence for dates: "9 April 1976." For months in tabular listings Synonymy in zoology must use the short form (taxon, author, or data sections, use three-letter abbreviations with no periods: "Jan, yearpage), with full reference at the end of the paper under Mar, Jun," etc. Omit space between initials of a personal name: "J.B. "Literature Cited." For botany, the long form (taxon, author, Jones." abbreviated journal or book title, volume, page, year, with no reference in "Literature Cited") is optional. Arrange and paginate sequentially every sheet of manuscript Text-reference system (author, yearpage used within the text, in the following order: (1) title page, (2) abstract, (3) contents, (4) with full citation in "Literature Cited" at the end of the text) must be foreword and/or preface, (5) text, (6) appendices, (7) notes section, used in place of bibliographic footnotes in all Contributions Series (8) glossary, (9) bibliography, (10) legends, (11) tables. Index copy and is strongly recommended in the Studies Series: "(Jones, may be submitted at page proof stage, but plans for an index should 1910:122)" or "...Jones (1910:122)." If bibliographic footnotes are be indicated when the manuscript is submitted. i